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A systematic revision of the South African Pettalidae (Arachnida : Opiliones : Cyphophthalmi) based on a combined analysis of discrete and continuous morphological characters with the description of seven new species Benjamin L. de Bivort A,B,C and Gonzalo Giribet B A Rowland Institute at Harvard, Harvard University, 100 Edwin Land Boulevard, Cambridge, MA 02142, USA. B Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA. C Corresponding author. Email: [email protected] Abstract. The cyphophthalmid family Pettalidae in South Africa is revised and seven new species are described from museum material collected between 1939 and 1985. Two of these are placed in the genus Purcellia and five in Parapurcellia, bringing the total number of described South African cyphophthalmids to 15. In addition, Purcellia peregrinator is transferred to the genus Parapurcellia. Phylogenetic analyses of discrete morphological and continuous morphometric characters, both separately and in combination, support the generic assignments and contribute towards a more detailed understanding of the systematics of the group in South Africa. In order to assess the stability of our phylogenetic results, the different morphological datasets were analysed under equal and implied weighting, as well as under several weighting schemes that varied the respective contribution to tree length of the discrete and continuous data partitions. These variations generated two phylogenetic hypotheses: (1) monophyly of the South African pettalids + Austropurcellia from north-eastern Australia as a derived clade within Pettalidae; and (2) polyphyly of the South African pettalids with Parapurcellia basal within Pettalidae. The latter hypothesis is congruent with previous molecular phylogenies of Cyphophthalmi, and has moderate bootstrap support. The sisterhood of Purcellia griswoldi, sp. nov. and P. lawrencei, sp. nov. receives high nodal support across analytic methods. New combination: Parapurcellia peregrinator (Lawrence, 1963). Additional keywords: Afromontane forests, Gondwanan biogeography, morphometrics, South Africa. Introduction The opilionid order Cyphophthalmi (often referred to as mite harvestmen) is represented by 17 species in Africa (Giribet 2000; Giribet and Prieto 2003). These include members in the tropical families Neogoveidae (three described species in the genus Parogovia Hansen, 1921), represented in western equatorial and western sub-Saharan Africa (Sierra Leone, Ivory Coast, Cameroon, Equatorial Guinea and Gabon; Hansen 1921; Juberthie 1969; Legg 1990; Giribet 2000; authors’ unpubl. data), and Ogoveidae (three described species in the genus Ogovea Roewer, 1923), represented in western equatorial Africa (Cameroon, Equatorial Guinea and Gabon; Hansen and Sørensen 1904; Hansen 1921; Giribet 2000; Giribet and Prieto 2003), plus a species of uncertain affinity from Kenya, Marwe coarctata Shear, 1985 (Shear 1985; Giribet and Boyer 2002). Within the territories of the former temperate Gondwana, the family Pettalidae is represented in South Africa by three genera, the monotypic Speleosiro Lawrence, 1931, and seven species in the genera Purcellia Hansen & Sørensen, 1904 and Parapurcellia Rosas Costa, 1950, plus a species in the monotypic genus Manangotria Shear & Gruber, 1996 in Madagascar. A second monotypic genus, Ankaratra Shear & Gruber, 1996, remains untested phylogenetically and is so far considered incertae sedis (Shear and Gruber 1996; Giribet 2000). Pettalidae contains eleven genera distributed across the continental fragments of temperate Gondwana (Boyer and Giribet 2007; Giribet and Boyer 2007): Chileogovea Roewer, 1961 (Chile) – Aoraki Boyer & Giribet, 2007, Rakaia Hirst, 1925 and Neopurcellia Forster, 1948 (New Zealand) – Austropurcellia Juberthie, 1988 (Queensland, Australia) – Karripurcellia Giribet, 2003 (Western Australia) – Pettalus Thorell, 1876 (Sri Lanka) – and the above mentioned African genera. Each genus is restricted to a single landmass, despite prior hypotheses of trans-Tasman distributions in Rakaia and Neopurcellia reported from New Zealand and Australia (see Boyer and Giribet 2007). There is considerable molecular and morphological support for the monophyly of many family-level clades within the Cyphophthalmi (Giribet and Boyer 2002; Boyer et al. 2007a), but the intergeneric relationships, especially within the family Pettalidae, are less resolved (e.g. Giribet 2003; Boyer and Giribet Ó CSIRO 13 December 2010 10.1071/IS10015 1445-5226/10/040371 CSIRO PUBLISHING www.publish.csiro.au/journals/is Invertebrate Systematics, 2010, 24, 371–406

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Page 1: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

A systematic revision of the South African Pettalidae(Arachnida Opiliones Cyphophthalmi) based on a combinedanalysis of discrete and continuous morphological characterswith the description of seven new species

Benjamin L de BivortABC and Gonzalo GiribetB

ARowland Institute at Harvard Harvard University 100 Edwin Land Boulevard Cambridge MA 02142 USABMuseum of Comparative Zoology and Department of Organismic and Evolutionary BiologyHarvard University 26 Oxford Street Cambridge MA 02138 USA

CCorresponding author Email debivortrowlandharvardedu

Abstract The cyphophthalmid family Pettalidae in South Africa is revised and seven new species are described frommuseummaterial collected between 1939 and 1985 Two of these are placed in the genusPurcellia andfive inParapurcelliabringing the total number of described South African cyphophthalmids to 15 In addition Purcellia peregrinator istransferred to the genus Parapurcellia Phylogenetic analyses of discrete morphological and continuous morphometriccharacters both separately and in combination support the generic assignments and contribute towards a more detailedunderstanding of the systematics of the group in South Africa In order to assess the stability of our phylogenetic results thedifferent morphological datasets were analysed under equal and implied weighting as well as under several weightingschemes that varied the respective contribution to tree length of the discrete and continuous data partitions These variationsgenerated two phylogenetic hypotheses (1) monophyly of the South African pettalids +Austropurcellia from north-easternAustralia as a derived clade within Pettalidae and (2) polyphyly of the South African pettalids with Parapurcellia basalwithin Pettalidae The latter hypothesis is congruent with previous molecular phylogenies of Cyphophthalmi and hasmoderate bootstrap support The sisterhood of Purcellia griswoldi sp nov and P lawrencei sp nov receives high nodalsupport across analytic methods New combination Parapurcellia peregrinator (Lawrence 1963)

Additional keywords Afromontane forests Gondwanan biogeography morphometrics South Africa

Introduction

The opilionid order Cyphophthalmi (often referred to as miteharvestmen) is represented by 17 species in Africa (Giribet2000 Giribet and Prieto 2003) These include members inthe tropical families Neogoveidae (three described species inthe genus Parogovia Hansen 1921) represented in westernequatorial and western sub-Saharan Africa (Sierra LeoneIvory Coast Cameroon Equatorial Guinea and GabonHansen 1921 Juberthie 1969 Legg 1990 Giribet 2000authorsrsquo unpubl data) and Ogoveidae (three described speciesin the genus Ogovea Roewer 1923) represented in westernequatorial Africa (Cameroon Equatorial Guinea and GabonHansen and Soslashrensen 1904 Hansen 1921 Giribet 2000Giribet and Prieto 2003) plus a species of uncertain affinityfrom KenyaMarwe coarctata Shear 1985 (Shear 1985 Giribetand Boyer 2002) Within the territories of the former temperateGondwana the family Pettalidae is represented in South Africaby three genera the monotypic Speleosiro Lawrence 1931 andseven species in the genera Purcellia Hansen amp Soslashrensen 1904and Parapurcellia Rosas Costa 1950 plus a species in the

monotypic genus Manangotria Shear amp Gruber 1996 inMadagascar A second monotypic genus Ankaratra Shear ampGruber 1996 remains untested phylogenetically and is so farconsidered incertae sedis (Shear andGruber 1996 Giribet 2000)

Pettalidae contains eleven genera distributed across thecontinental fragments of temperate Gondwana (Boyer andGiribet 2007 Giribet and Boyer 2007) Chileogovea Roewer1961 (Chile) ndashAorakiBoyerampGiribet 2007RakaiaHirst 1925andNeopurcellia Forster 1948 (NewZealand) ndashAustropurcelliaJuberthie 1988 (QueenslandAustralia) ndashKarripurcelliaGiribet2003 (Western Australia) ndash Pettalus Thorell 1876 (Sri Lanka) ndashand the abovementionedAfrican genera Each genus is restrictedto a single landmass despite prior hypotheses of trans-Tasmandistributions in Rakaia and Neopurcellia reported from NewZealand and Australia (see Boyer and Giribet 2007)

There is considerable molecular and morphological supportfor the monophyly of many family-level clades within theCyphophthalmi (Giribet and Boyer 2002 Boyer et al 2007a)but the intergeneric relationships especially within the familyPettalidae are less resolved (eg Giribet 2003 Boyer and Giribet

CSIRO 13 December 2010 101071IS10015 1445-522610040371

CSIRO PUBLISHING

wwwpublishcsiroaujournalsis Invertebrate Systematics 2010 24 371ndash406

2007 2009 de Bivort et al 2010) Molecular evidence favoursthe monophyly of all genera as currently defined although only afew (Aoraki Rakaia Pettalus) have been thoroughly sampled inmolecular studies (Boyer and Giribet 2007 Boyer et al 2007a2007b) In these studies the African pettalids were restricted torepresentatives of only three species one in the genus Purcelliaand two in the genus Parapurcellia and South Africa wasparaphyletic with respect to other landmasses

The described South African pettalid fauna is currentlydistributed in two geographic regions (Fig 1) Two speciesare found in the Western Cape province specifically in andaround Table Mountain Purcellia illustrans Hansen ampSoslashrensen 1904 has been collected in forests surroundingTable Mountain Speleosiro argasiformis Lawrence 1931 isa large (5mm) cyphophthalmid found in the Wynberg cavesystem in Table Mountain (Juberthie 1970) The formerspecies is a medium-sized cyphophthalmid with morphologicalcharacteristics typical of other members of South Africanpettalids such as relatively unmodified opisthosomal posteriortergites in males and females second palp articles that are lessthan half ornamented and bilobed anal plates in males Forillustrations of these and other cyphophthalmid morphologicalterms see de Bivort et al (2010) One conspicuous featureof P illustrans is the position of its anal plate which is shiftedanteriorly onto the ventral surface of the animal comparedwith other pettalids Speleosiro argasiformis has conspicuoustroglomorphic modifications such as elongated appendages

While Speleosiro is monotypic Purcellia is currentlyrepresented by two additional species found in eastern SouthAfrica Purcellia transvaalica Lawrence 1963 is found inthe Hanglip forest of the Limpopo Province and has anappearance typical of other Purcellia species Purcelliaperegrinator Lawrence 1963 found in Mariepskop ForestMpumalanga Province has characteristics more typical ofthe genus Parapurcellia which has a distribution in the NatalDrakensberg and NatalndashZululand Coast of the Eastern Cape and

KwazulundashNatal Provinces In fact Lawrence (1963 279) statedthat lsquoThis species resembles [Parapurcellia] silvicola fromZululand in the shape of the tarsus IVrsquo In particular it hasozophores oriented at 45 off the lateral margin rather thandorsal ozophores projections of the anterior gonostome walla small (~25mm) body length and no evidence of functionaleyes under light microscopy ndash unlike Purcellia which haslensless eyes situated in the bases of the ozophores Based onthese characters as well as several others discussed belowand consistent placement of Purcellia peregrinator withinParapurcellia in our phylogenetic analyses we reassignPurcellia peregrinator to Parapurcellia establishing the newcombination Parapurcellia peregrinator (Lawrence 1963)leaving the number of described species in Parapurcellia atfive Parapurcellia fissa (Lawrence 1939) and Parapurcelliasilvicola (Lawrence 1939) are distinctive due to their deeplylobed posterior tergites In the remaining two speciesParapurcellia monticola (Lawrence 1939) and Parapurcelliarumpiana (Lawrence 1933) the posterior tergites are relativelyunmodified and their morphologies are typical of small pettalidsalthough the former species is distinguishable from the otherSouth African species by its tall profile

The Natal Museum of South Africa (NMSA) kindlyprovided us with undescribed cyphophthalmid specimenscollected by several arachnologists between 1939 and 1985We identified two distinct morphospecies collected in theKnysna forest that straddles the border of the Eastern andWestern Cape Provinces which appear to belong in PurcelliaAn additional four morphospecies from localities in the EasternCape and KwazulundashNatal Provinces segregated years ago byW Starega who assigned tentative names to some of the speciesappear to belong in Parapurcellia We have chosen to use hisnames as much as possible to honour his early recognition of thedifferent morphologies These are described as new speciesbelow and our placement of them into their respective generais supported by morphological phylogenetic analysis

Speleosiro argasiformisPurcellia illustrans

Parapurcellia peregrinator

Parapurcellia fissa

Purcellia griswoldi sp nov

Parapurcellia amatola sp nov

Parapurcellia staregai sp nov

Parapurcellia minuta sp novParapurcellia natalia sp nov

Purcellia lawrencei sp nov

Parapurcellia monticolaParapurcellia rumpiana

Parapurcellia silvicola

Purcellia transvaalica

NORTHERN CAPE

WESTERN CAPE

EASTERN CAPE

FREE STATE

NORTH WEST

LIMPOPO

GAU-TENG

KWAZULU-NATAL

MPUMA-LANGA

NORTHERN CAPE

WESTERN CAPE

EASTERN CAPE

FREE STATE

NORTH WEST

LIMPOPO

GAU-TENG

KWAZULU-NATAL

MPUMA-LANGA

Parapurcellia convexa sp nov

Fig 1 Mapofpettalid type collection localities inSouthAfricaCircles indicate localities of speciesdescribedbeforethiswork Stars indicate new species described herein Square indicatesParapurcellia peregrinator (Lawrence 1963)new comb whose genus is reassigned herein

372 Invertebrate Systematics B L de Bivort and G Giribet

The NMSA collections along with most collections ofpreviously described species were made before the era ofmolecular phylogenetics Their long-term storage in 70ethanol has precluded the possibility of analysing the species-level relationships of the South African pettalids using molecular

sequences In order to investigate the phylogenetic relationshipsof the SouthAfricanmaterial we chose to perform aphylogeneticanalysis under the parsimony criterion using all availablemorphological data Attempts at lsquototal evidencersquo analysis haveproved successful in past analyses of the Cyphophthalmi

Table 1 Discrete character matrixCharacter codings are explained in Appendix 2

Taxa Characters000000000 1111111111 2222222222 3333333333 4444444444 5555555555 666

123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012

Parapurcellia staregai sp nov 032200011 031101100 011001100 00-000100 0112000100 1130100100 0

Parapurcellia amatolae sp nov 032200011 031100100 1101011000 00-0020100 0112000100 1130100100 0

Parapurcellia convexa sp nov 032200011 031100100 0101011100 00-0020100 0112000110 1120100101 0

Parapurcellia natalia sp nov 032200011 031000100 011101100 00-000100 0112000110 1130100100 0

Parapurcellia minuta sp nov 032200011 031101100 0101011000 00-0020110 0112000110 10-0100100 0

Parapurcellia peregrinator 032200011 021100100 111101110 00-00300 0112000110 1120100100 0

Parapurcellia silvicola 032200011 031100000 0101011100 00-0020100 01100110 0100100101 0

Parapurcellia monticola 032200011 021100000 0101011100 00-0020100 01100110 1130100100 0

Parapurcellia rumpiana 032200011 021001100 010111101 0-30100 01100110 110100100 0

Parapurcellia fissa 032200011 031100100 0101110 30100 0100110 130100100 0

Purcellia transvaalica 222200011 020101100 110101100 00-0020 0112100100 1140110100 0

Purcellia illustrans 222200011 000100000 1101011101 00-0020100 0112100100 0140110100 001

Purcellia griswoldi sp nov 222200010 020100100 1001011100 00-0020100 0112100100 1140110100 011

Purcellia lawrencei sp nov 222200011 030100100 110101100 00-0020100 0112100100 1140110100 0

Speleosiro argasiformis 022200011 000100000 1100011101 0-0020100 0111000100 0140110100 0

Rakaia isolata 232200011 000101000 110101100 00-0020000 0100110 0100101101 0

Rakaia lindsayi 232200011 000100000 1100010210 00-0020000 01100110 0100100101 0

Rakaia magna australis 232200011 000101000 1100011200 00-0020000 01100110 0100100100 0

Rakaia media 232200011 000100000 1101011200 00-0020000 011200110 0100100100 0

Rakaia solitaria 132200011 000100000 1101011200 00-0020000 01100110 0100101100 0

Rakaia sorenseni digitata 232200011 000100000 1101011100 00-0020050 010ndashndash00110 00-01-1000 0

Aoraki crypta 232200011 000100000 1110010101 00-0020000 011200110 01311-0000 0

Aoraki denticula 232200011 000100000 1110010201 00-0021000 011200110 01301-0001 0

Aoraki inerma 232200011 000100000 1110010201 00-0020000 011200110 01311-0000 0

Aoraki longitarsa 232200011 000100000 1110010201 00-0021000 0100110 0100100101 1

Aoraki tumidata 232200011 000101000 11001011 0-2 112100111 10-0101101 0

Karripurcellia harveyi 232200011 000100010 1110010210 00-0020000 010ndashndash10100 00-00-0000 0

Karripurcellia peckorum 232200011 000101010 1110010210 00-0020000 010ndashndash10100 00-00-0000 001

Karripurcellia sierwaldae 232200011 000101010 1100011200 00-0020000 010ndashndash10100 00-00-0000 0

Austropurcellia scoparia 232200011 000100000 1101011101 00-0030100 0114000110 01101-0000 11

Austropurcellia woodwardi 232200011 020100000 111101 00-0030100 011000110 0130100100 1

Chileogovea jocasta 132200011 001100000 1110010101 00-1140000 0112100100 00-00-0000 0

Chileogovea oedipus 132100011 001100000 1110010211 00-1141000 0112100101 10-00-0000 01

Pettalus lampetides 122200011 001100010 1110110001 00-0041001 010ndashndash02100 00-01-1000 01

Pettalus cimiciformis 122100011 00310001 1110010101 00-4100 010ndashndash02100 00-01-1000 0

Troglosiro aelleni 010210011 001000000 2110000201 0110030000 001003-00 00-00-0000 0

Troglosiro longifossa 010210011 0010100100 2110000101 0110030000 001003-00 00-00-0010 0

Troglosiro ninqua 010210011 0010100000 2110000000 0110030000 001003-00 00-00-0010 0

Huitaca ventralis 010200101 111-100010 2110010201 0110030001 001003-00 00-00-0000 00

Neogovea sp 010200111 1310101010 1101000 0 003-00 00-00-0000 0

Metagovea philipi 010200111 12110000 2110000101 0110030001 001003-00 00-00-0000 001

Cyphophthalmus duricorius 010211011 0020000101 0000010100 00-0010000 001103-01 10-00-0000 011

Parasiro coiffaiti 000201010 0010000101 0100010000 0111120000 000ndashndash01000 00-00-0000 010

Siro acaroides 010211010 0010000100 0000000000 00-1110000 0011103-00 10-01-0000 011

Siro rubens 010211011 0020000100 0000020100 00-0010000 0011103-01 00-00-0000 011

Suzukielus sauteri 012200011 001000100 0000020100 1101120100 0013000101 00-00-0000 011

Stylocellus ramblae 111200101 000010001 011011021 00-0151010 020ndashndash00000 00-00-0000 0

Stylocellus tambusisi 111200100 121010001 01100102 00-001010 020ndashndash00000 10-00-0000 0

Fangensis cavernarus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001

Fangensis spelaeus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001

Systematic revision of South African pettalids Invertebrate Systematics 373

Tab

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Con

tinu

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itap

pearsin

the11

weigh

tedcombinedan

alysis

Character

definitio

nsaregivenin

App

endix3

12

34

56

78

910

1112

1314

1516

1718

1920

2122

2324

2526

2728

2930

3132

3334

3536

3738

39

Parap

urcelliastaregaisp

nov

054

059

058

052

053

053

059

047

064

051

058

063

047

067

069

075

048

071

070

056

055

049

062

073

057

057

052

052

054

066

054

068

048

048

056

056

067

061

052

Parap

urcelliaam

atolaesp

nov

047

061

057

056

056

064

052

047

065

057

053

072

051

062

061

059

062

070

066

056

042

063

062

053

053

058

058

057

072

055

050

061

043

049

056

056

066

062

049

Parapurcelliaconvexa

spn

ov

048

069

058

075

067

066

049

053

060

051

050

078

081

063

074

051

064

062

071

059

056

065

057

059

045

066

085

075

071

070

068

052

057

051

056

056

061

060

054

Parapurcellianatalia

sp

nov

044

056

057

052

053

057

049

047

057

049

060

063

048

050

071

061

105

082

074

060

053

059

061

076

059

059

052

052

065

056

043

065

043

047

056

056

071

058

049

Parapurcelliaminuta

spn

ov

042

056

056

052

053

046

050

047

057

054

081

075

047

068

072

082

048

069

062

060

059

069

059

050

052

057

052

052

043

067

055

064

057

052

056

056

067

058

048

Parapurcelliaperegrinator

059

061

058

059

055

057

059

047

077

055

065

065

048

063

049

048

066

061

057

061

055

056

057

062

071

059

060

055

058

061

053

062

053

047

056

056

065

059

059

Parapurcelliasilvicola

055

063

063

076

064

049

062

050

062

062

060

072

077

053

079

050

065

079

070

059

060

065

064

064

057

071

080

067

047

097

059

068

040

059

056

056

064

058

070

Parapurcelliamonticola

054

061

059

061

053

056

045

053

073

052

058

074

045

055

064

073

050

070

059

057

065

062

058

064

061

063

063

052

058

054

066

049

074

051

056

056

066

063

062

Parap

urcelliarumpiana

051

059

056

057

053

050

040

053

067

078

046

073

075

057

073

080

037

074

071

057

061

058

054

056

059

059

060

052

049

062

066

039

061

046

056

056

075

060

055

Parapurcelliafissa

043

064

064

052

053

056

059

053

065

052

055

070

071

069

076

063

055

066

060

057

063

079

078

060

051

085

052

052

062

036

068

083

053

053

056

056

068

060

052

Purcelliatransvaalica

069

064

057

052

053

048

060

058

061

058

041

071

070

057

074

050

061

053

072

050

067

050

056

054

056

056

052

052

046

056

054

051

050

036

056

056

074

058

048

Purcelliaillustrans

072

063

055

052

053

052

060

087

081

056

061

063

062

061

077

049

059

075

071

051

065

058

051

078

087

054

052

052

050

089

096

053

076

036

056

056

078

061

042

Purcelliagriswoldisp

nov

062

063

057

057

053

043

062

055

067

053

065

066

062

044

064

068

050

057

058

048

075

059

055

053

058

058

057

052

041

061

061

060

060

040

056

056

075

057

045

Purcellialawrenceisp

nov

062

061

056

057

053

051

066

057

057

053

059

069

067

047

054

066

053

068

067

048

080

063

052

056

050

055

057

052

050

063

068

064

068

039

056

056

075

053

044

Speleosiro

argasiform

is097

077

060

075

059

073

091

048

071

051

049

044

063

039

056

055

068

031

035

045

108

046

054

047

050

057

064

056

061

053

056

060

057

047

056

056

070

058

043

Rakaiaisolate

059

077

067

102

058

069

066

061

059

063

067

077

077

070

062

046

077

054

068

052

062

062

072

061

043

076

104

058

071

066

061

068

059

062

056

056

058

064

099

Rakaialin

dsayi

061

063

062

083

053

059

053

057

054

066

071

056

071

077

067

086

046

079

059

051

057

071

058

057

061

070

082

052

061

063

037

055

037

056

056

056

062

066

062

Rakaiamag

naau

stralis

074

068

064

078

061

085

070

047

077

085

072

052

071

057

043

053

067

068

062

054

062

062

066

068

058

060

072

059

078

065

065

059

067

053

056

056

065

058

075

Rakaiamedia

062

070

062

063

057

055

077

062

066

052

066

058

070

081

066

087

041

065

064

049

057

057

068

058

068

061

062

057

054

058

054

077

057

064

056

056

061

063

077

Rakaiasolitaria

057

065

059

071

053

065

052

060

079

053

074

080

051

055

051

046

070

065

068

053

077

043

055

037

060

065

073

052

069

069

067

055

065

066

056

056

053

062

074

Rakaiasorensenidigitata

057

071

055

054

055

075

071

080

065

066

056

093

052

085

038

054

064

069

065

051

075

075

053

066

069

054

055

056

083

060

059

080

058

074

056

056

047

053

078

Aorakicrypta

071

079

061

058

059

085

069

088

041

059

049

059

043

072

059

047

071

060

064

052

060

037

051

039

053

071

057

058

081

067

062

061

053

039

056

056

077

075

086

Aorakidenticula

057

067

055

068

062

063

048

080

054

054

056

065

051

085

058

060

067

068

071

053

066

065

051

057

069

055

071

064

066

059

063

050

063

044

075

072

071

070

070

Aorakiinerma

067

066

057

063

063

062

056

070

075

056

044

066

065

077

053

051

066

060

059

054

073

066

053

053

063

057

062

063

060

054

060

050

059

062

056

056

059

072

074

Aorakilong

itarsa

066

076

073

098

109

065

058

069

056

058

083

056

049

062

028

049

061

063

064

050

061

059

066

079

060

091

094

107

064

076

062

055

058

064

056

056

059

129

062

Aorakitumidata

064

072

058

081

069

079

070

072

063

068

071

073

079

083

054

080

060

073

070

050

062

072

055

054

062

060

083

070

081

085

062

070

050

070

075

096

080

059

071

Karripurcellia

harveyi

067

052

058

061

066

051

079

089

062

052

069

056

073

060

074

075

037

050

043

055

056

063

060

069

091

052

060

065

049

066

050

072

048

067

056

056

047

057

058

Karripurcellia

peckorum

068

062

057

071

068

069

062

070

051

094

057

064

070

057

052

045

071

046

037

055

060

064

057

059

076

054

069

067

064

070

046

053

037

068

056

056

047

056

056

Karripurcellia

sierwaldae

056

056

057

052

053

042

062

061

067

088

056

061

076

063

070

057

060

043

032

055

062

069

062

067

068

053

052

052

039

067

057

064

051

073

064

087

047

057

060

Austropurcelliascoparia

055

050

054

052

053

067

052

068

054

051

045

065

050

064

057

044

071

065

074

059

071

072

051

054

084

053

052

052

071

052

082

055

088

062

056

056

061

091

053

Austrop

urcelliawoo

dwardi

054

065

058

052

058

056

061

069

053

053

042

066

049

066

068

053

057

062

068

054

063

047

049

036

050

067

052

060

056

065

079

064

068

059

056

056

047

061

053

Chileogovea

jocasta

049

053

055

052

054

058

050

057

055

049

047

049

058

060

055

063

050

050

058

051

074

055

053

053

060

053

052

055

061

057

069

057

068

066

082

088

082

053

069

Chileog

ovea

Oedipus

070

053

054

052

053

064

070

074

057

062

053

052

043

064

054

051

058

072

078

053

071

012

049

043

054

052

052

052

061

052

063

061

070

062

092

091

083

051

048

Pettaluslampetid

es063

066

078

052

092

067

057

058

053

055

063

039

048

055

057

063

064

068

068

046

064

050

095

090

044

084

052

093

065

042

050

052

066

074

056

056

047

052

055

Pettaluscimicifo

rmis

100

110

140

066

095

089

082

076

066

058

074

033

039

021

057

058

058

056

046

041

056

085

125

064

044

116

059

074

070

039

047

055

071

068

056

056

047

050

061

Troglosiroaelleni

059

053

059

054

066

066

064

057

054

053

052

040

067

066

072

060

064

044

053

076

045

064

066

075

062

054

054

067

068

054

061

066

064

074

056

056

047

055

058

Troglosirolong

ifossa

051

045

055

052

073

064

048

070

049

060

057

042

057

053

069

043

069

053

064

079

041

059

055

061

066

052

052

081

070

049

049

052

055

074

056

056

047

056

053

Troglosironinq

ua059

045

057

053

086

068

049

074

049

066

048

044

074

054

064

052

063

048

059

076

044

054

063

072

068

052

054

092

069

062

076

048

070

074

056

056

047

057

072

Huitaca

ventralis

075

045

057

052

060

036

084

047

105

094

063

054

058

067

072

068

054

040

056

075

057

046

058

070

053

052

052

058

039

045

038

057

047

074

056

056

047

052

048

Neogo

veasp

067

056

058

065

070

031

070

075

081

099

042

055

067

047

063

068

042

031

045

074

050

038

061

073

061

052

063

069

033

035

035

055

044

074

056

056

047

050

045

Metagovea

philipi

045

045

054

052

053

041

045

057

065

055

056

056

066

051

053

054

056

048

062

078

057

057

053

050

031

052

052

052

036

042

056

048

073

074

056

056

047

053

068

Cypho

phthalmus

duricorius

050

045

055

052

053

058

041

050

039

060

067

044

044

042

027

050

072

058

065

078

039

065

057

071

065

052

052

052

062

059

067

044

072

074

056

056

047

056

065

Parasirocoiffaiti

044

045

055

054

055

058

056

047

050

052

079

050

041

060

056

080

051

064

061

086

042

064

060

062

058

052

056

057

068

053

052

081

063

068

056

056

055

053

064

Siro

acaroids

046

055

054

052

053

055

053

049

044

055

047

051

067

060

070

051

057

058

071

069

048

068

053

044

061

053

052

052

058

062

061

062

055

051

056

056

071

055

068

Siro

rubens

049

045

056

052

053

042

077

050

041

055

065

039

057

053

062

082

037

047

052

072

035

073

059

052

086

052

052

052

036

054

085

102

100

066

119

102

056

050

049

Suzukielus

sauteri

059

045

056

052

053

060

062

048

050

051

063

068

069

065

064

066

056

082

071

075

048

071

056

067

057

052

052

052

062

079

072

066

063

071

098

091

047

056

061

Stylocellusramblae

052

045

055

052

053

064

050

060

052

048

083

058

047

056

053

055

076

045

046

092

068

060

054

046

048

052

052

052

067

053

058

053

074

074

056

056

047

053

060

Stylocellustambusisi

074

045

057

052

053

066

049

059

046

067

058

052

050

041

029

059

076

047

056

082

066

060

057

070

056

052

052

052

061

047

055

041

065

074

056

056

047

052

062

Fan

gensiscavernarus

081

064

065

052

053

073

046

047

049

065

080

054

076

053

065

048

079

048

026

075

061

073

065

078

081

059

052

052

066

063

049

038

051

074

056

056

047

053

048

Fan

gensisspelaeus

073

045

057

052

053

078

093

047

050

061

093

065

077

056

064

076

053

068

030

076

054

069

056

069

066

052

052

052

076

068

055

086

061

074

056

056

047

057

054

374 Invertebrate Systematics B L de Bivort and G Giribet

4041

4243

4445

4647

4849

5051

5253

5455

5657

5859

6061

6263

6465

6667

6869

7071

7273

7475

7677

78

Parap

urcelliastaregaisp

nov

054

049

048

053

061

054

046

048

065

057

060

056

057

059

069

055

074

063

048

049

055

058

069

062

068

080

057

066

053

057

056

052

068

059

066

054

070

051

062

Parap

urcelliaam

atolaesp

nov

047

058

055

044

066

054

046

050

081

058

068

069

065

064

068

055

068

055

049

032

047

064

064

070

066

059

053

073

046

043

035

075

066

053

068

048

065

062

056

Parapurcelliaconvexa

spn

ov

036

030

068

095

053

054

055

050

064

057

053

052

056

065

070

055

069

066

055

062

053

069

052

075

070

059

063

067

049

055

069

062

077

034

062

057

071

052

062

Parapurcellianatalia

sp

nov

050

046

050

051

054

054

046

046

079

060

056

066

072

063

064

055

062

060

057

058

046

068

053

063

063

069

042

063

045

054

073

050

062

073

056

054

060

048

054

Parap

urcelliaminuta

spn

ov

046

046

054

041

030

054

047

049

075

060

059

050

058

059

066

055

062

062

057

040

062

066

072

061

062

076

044

064

052

056

058

076

095

069

053

058

062

057

054

Parap

urcelliaperegrinator

058

058

058

050

049

054

046

054

071

063

049

056

070

062

060

055

062

060

057

070

053

063

073

077

079

056

052

078

049

061

058

057

064

079

052

048

066

055

055

Parapurcelliasilvicola

052

054

062

050

044

054

050

048

064

058

063

059

055

062

069

055

064

061

052

051

062

067

072

070

066

019

045

064

059

059

048

061

074

066

065

073

070

064

057

Parapurcelliamonticola

067

073

086

060

055

054

053

051

066

057

050

062

071

063

070

055

060

065

058

053

044

064

061

039

072

075

047

080

048

061

056

047

066

061

066

047

061

053

056

Parap

urcelliarumpian

a040

035

072

081

058

054

055

045

075

057

048

067

074

063

065

055

082

059

056

072

052

066

051

064

073

068

049

076

073

066

056

058

070

035

054

090

078

093

058

Parapurcelliafissa

060

057

065

048

059

054

057

060

063

057

049

056

066

063

060

055

062

061

051

064

056

065

062

038

074

051

057

079

050

059

043

048

069

061

067

051

071

055

060

Purcelliatransvaalica

064

068

060

051

053

054

056

057

062

055

044

054

065

055

069

055

077

068

052

054

044

068

069

066

064

058

060

058

060

071

050

043

054

051

073

055

054

056

058

Purcelliaillustrans

111

080

101

074

037

054

082

067

066

052

047

046

060

066

057

055

056

056

055

047

052

066

063

057

066

048

055

077

052

068

051

043

059

048

053

047

072

054

059

Purcelliagriswoldisp

nov

067

078

063

048

049

054

054

049

065

057

045

050

066

061

065

055

053

061

059

046

050

063

074

056

070

071

070

066

058

070

035

050

051

069

049

058

077

052

056

Purcellialawrenceisp

nov

071

081

067

052

046

054

056

052

089

054

046

040

054

061

066

055

059

059

069

047

038

074

067

054

072

068

056

056

056

065

051

050

050

050

042

062

068

052

057

Speleosiro

argasiform

is057

056

054

045

060

054

047

044

055

066

041

058

078

061

045

055

058

061

057

044

076

049

090

079

055

054

085

058

077

089

050

052

051

070

043

041

050

048

060

Rakaiaisolate

066

075

061

060

062

054

061

079

032

073

059

069

071

063

059

055

066

060

066

044

061

052

039

053

069

056

038

058

049

050

049

077

071

058

077

062

065

064

059

Rakaialin

dsayi

050

060

058

056

068

054

057

054

070

057

067

065

060

067

060

055

055

056

068

059

073

059

060

044

078

074

049

053

060

059

050

064

057

085

059

055

064

066

058

Rakaiamag

naau

stralis

060

047

055

055

055

054

046

052

059

078

064

074

072

066

054

055

061

064

073

060

060

064

050

056

070

047

058

068

062

052

053

074

049

049

058

061

070

077

058

Rakaiamedia

060

082

041

052

062

054

061

052

065

078

064

082

075

061

058

055

060

062

075

060

047

059

043

059

063

071

060

064

049

037

076

056

070

050

061

063

070

051

060

Rakaiasolitaria

065

071

056

052

051

054

060

066

068

067

074

075

063

069

053

055

078

056

065

060

053

064

047

055

063

051

057

081

050

048

074

072

056

065

075

069

073

052

058

Rakaiasorensenidigitata

076

071

066

060

034

054

087

052

064

060

049

030

039

084

055

055

065

065

065

060

068

057

072

072

072

051

067

069

061

062

061

078

076

078

077

073

071

070

048

Aorakicrypta

071

069

053

060

054

054

082

048

068

067

068

063

051

069

055

055

059

073

071

054

064

047

075

078

071

056

080

040

068

049

067

081

042

064

068

041

047

057

048

Aorakidenticula

075

068

054

061

068

054

085

055

052

057

068

055

048

077

054

055

070

060

056

065

047

073

071

049

074

066

053

055

057

035

085

061

070

067

058

040

059

054

060

Aorakiinerma

069

070

071

078

065

054

069

046

068

060

064

063

052

072

050

055

081

064

054

050

058

060

062

072

042

055

057

073

070

068

054

052

047

043

079

046

057

055

061

Aorakilong

itarsa

069

066

057

062

073

054

067

078

064

059

075

063

056

071

056

055

065

073

057

069

073

043

074

071

064

050

071

054

064

049

071

078

050

052

062

056

063

057

062

Aorakitumidata

066

062

067

047

063

054

073

062

066

055

074

087

071

070

065

055

051

065

072

062

095

023

074

082

079

081

077

063

082

060

064

105

068

085

055

059

071

069

069

Karripurcellia

harveyi

050

061

072

074

086

054

084

076

057

054

047

047

064

062

057

055

056

054

061

077

075

053

060

073

053

047

066

059

064

070

054

051

055

061

045

052

054

050

060

Karripurcellia

peckorum

046

050

066

077

087

054

067

056

050

067

045

047

062

064

056

055

059

059

074

071

069

056

061

059

063

045

055

049

082

067

054

055

042

054

081

071

051

090

055

Karripurcellia

sierwaldae

050

052

064

067

068

054

062

061

060

050

060

044

044

064

065

055

052

060

051

060

060

058

057

064

061

060

063

070

079

074

051

040

045

058

056

081

057

097

048

Austropurcelliascoparia

079

078

069

066

053

054

074

064

051

052

080

056

037

089

038

055

056

057

080

050

057

076

067

062

060

048

053

056

052

062

064

060

061

068

062

066

062

052

063

Austrop

urcelliawoo

dwardi

072

067

074

061

054

054

075

079

049

062

075

067

047

074

036

055

058

058

053

054

051

086

061

052

056

059

038

054

054

065

060

046

058

049

063

080

067

054

048

Chileogovea

jocasta

067

064

065

054

054

054

061

052

054

055

069

066

052

069

045

055

049

058

057

057

063

066

042

053

059

068

068

059

056

074

054

056

057

060

043

057

046

050

048

Chileog

ovea

oedipu

s064

068

070

070

048

054

070

076

081

056

070

056

046

069

056

055

054

059

054

085

072

060

047

044

076

057

066

055

072

074

048

064

054

076

046

052

041

060

048

Pettaluslampetid

es063

055

059

058

058

054

057

051

063

063

053

077

084

052

075

055

058

051

057

058

054

052

051

057

060

063

062

052

073

067

054

051

039

075

083

040

034

053

061

Pettaluscimicifo

rmis

063

061

050

062

048

062

060

058

044

065

044

054

079

048

070

055

042

048

069

049

060

051

057

051

058

062

082

054

051

061

077

059

060

053

072

046

062

051

057

Troglosiroaelleni

059

056

029

089

066

085

058

054

066

046

078

071

050

062

045

090

056

060

068

076

055

055

066

058

045

064

076

054

064

065

059

049

055

055

046

049

037

052

060

Troglosirolongifo

ssa

059

055

058

070

077

085

077

053

061

045

069

071

058

048

054

093

038

065

070

085

059

056

052

082

044

051

071

049

056

045

075

053

050

053

038

063

073

064

048

Troglosironinq

ua066

051

072

073

079

083

077

061

060

041

068

067

053

045

064

092

062

059

058

067

059

053

070

075

052

061

082

049

075

076

046

055

048

051

036

053

045

068

048

Huitaca

ventralis

045

045

040

050

059

085

046

058

056

041

049

070

083

032

075

081

031

044

049

051

062

061

070

030

046

062

071

090

108

096

085

062

043

073

043

063

033

084

048

Neogo

veasp

042

067

051

086

073

087

071

066

031

046

039

066

087

037

059

055

066

130

104

061

066

071

062

039

042

072

075

076

074

077

079

055

050

036

084

064

038

095

061

Metag

ovea

philipi

057

068

061

073

063

085

062

044

040

041

066

069

059

039

057

055

061

044

080

064

056

085

058

045

035

059

041

075

059

063

053

049

046

047

065

054

034

059

048

Cypho

phthalmus

duricorius

060

046

053

056

061

086

050

058

065

056

053

064

073

054

051

087

060

064

057

082

060

061

060

065

048

047

064

036

055

052

070

059

066

065

071

077

072

064

078

Parasirocoiffaiti

056

064

071

050

066

081

046

074

063

060

089

070

051

051

055

078

055

065

053

062

063

060

032

073

042

079

053

052

044

048

078

068

088

061

060

077

070

055

048

Siro

acaroides

052

040

038

053

081

085

049

077

058

060

057

079

073

055

059

096

032

061

049

068

045

071

040

057

055

059

058

043

065

064

063

065

066

050

059

067

056

059

048

Siro

rubens

063

050

064

053

063

085

050

069

056

049

076

069

054

060

051

087

073

067

078

089

056

064

034

060

046

084

064

038

056

054

068

064

076

049

045

063

068

058

073

Suzukielus

sauteri

064

047

063

048

073

054

047

101

069

053

063

050

048

052

075

055

060

062

067

077

059

058

042

069

068

069

052

050

049

048

075

061

066

056

069

065

074

050

068

Stylocellusramblae

060

072

062

053

058

054

062

073

041

098

074

042

040

035

071

055

025

044

038

053

105

053

072

051

030

061

082

052

056

052

067

075

059

072

063

066

048

047

099

Stylocellustambusisi

056

063

061

052

060

054

056

069

047

096

073

050

041

040

067

055

063

044

038

042

064

060

074

050

036

048

063

044

060

049

067

060

058

078

057

085

046

062

087

Fan

gensiscavernarus

047

044

040

056

075

054

046

070

036

085

050

042

058

041

084

055

077

044

038

066

076

035

052

064

049

055

070

046

051

044

056

067

062

071

050

058

060

057

096

Fan

gensisspelaeus

049

061

044

055

079

054

046

070

047

086

046

035

058

042

105

055

087

044

038

065

077

029

068

065

060

084

050

047

051

045

078

070

083

057

054

071

065

056

096

Systematic revision of South African pettalids Invertebrate Systematics 375

combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)

Methods and abbreviations

Specimens used in this study were provided by the followinginstitutions

AMNH American Museum of Natural History New York(USA)

BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard

University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand

Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo

Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg

(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg

Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)

Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg

(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen

Copenhagen (Denmark)

Specimen handling and illustration

In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was

photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy

Character coding

Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3

Phylogenetic analyses

The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)

The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the

376 Invertebrate Systematics B L de Bivort and G Giribet

continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data

Taxonomy

Order OPILIONES Sundevall

Suborder CYPHOPHTHALMI Simon

Family PETTALIDAE Simon

Genus Speleosiro Lawrence 1931

Diagnosis

Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII

Type species

Speleosiro argasiformis Lawrence 1931

Species included

Speleosiro argasiformis Lawrence 1931

Distribution

SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province

Remarks

This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement

Genus Purcellia Hansen amp Soslashrensen 1904

Diagnosis

Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)

Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)

Type species

Purcellia illustrans Hansen amp Soslashrensen 1904

Species included

Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov

Distribution

South Africa Limpopo Eastern Cape and Western CapeProvinces

Remarks

This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa

Purcellia griswoldi sp nov

(Figs 2 3 Table 3)

Material examined

Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985

Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981

Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981

Systematic revision of South African pettalids Invertebrate Systematics 377

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 2: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

2007 2009 de Bivort et al 2010) Molecular evidence favoursthe monophyly of all genera as currently defined although only afew (Aoraki Rakaia Pettalus) have been thoroughly sampled inmolecular studies (Boyer and Giribet 2007 Boyer et al 2007a2007b) In these studies the African pettalids were restricted torepresentatives of only three species one in the genus Purcelliaand two in the genus Parapurcellia and South Africa wasparaphyletic with respect to other landmasses

The described South African pettalid fauna is currentlydistributed in two geographic regions (Fig 1) Two speciesare found in the Western Cape province specifically in andaround Table Mountain Purcellia illustrans Hansen ampSoslashrensen 1904 has been collected in forests surroundingTable Mountain Speleosiro argasiformis Lawrence 1931 isa large (5mm) cyphophthalmid found in the Wynberg cavesystem in Table Mountain (Juberthie 1970) The formerspecies is a medium-sized cyphophthalmid with morphologicalcharacteristics typical of other members of South Africanpettalids such as relatively unmodified opisthosomal posteriortergites in males and females second palp articles that are lessthan half ornamented and bilobed anal plates in males Forillustrations of these and other cyphophthalmid morphologicalterms see de Bivort et al (2010) One conspicuous featureof P illustrans is the position of its anal plate which is shiftedanteriorly onto the ventral surface of the animal comparedwith other pettalids Speleosiro argasiformis has conspicuoustroglomorphic modifications such as elongated appendages

While Speleosiro is monotypic Purcellia is currentlyrepresented by two additional species found in eastern SouthAfrica Purcellia transvaalica Lawrence 1963 is found inthe Hanglip forest of the Limpopo Province and has anappearance typical of other Purcellia species Purcelliaperegrinator Lawrence 1963 found in Mariepskop ForestMpumalanga Province has characteristics more typical ofthe genus Parapurcellia which has a distribution in the NatalDrakensberg and NatalndashZululand Coast of the Eastern Cape and

KwazulundashNatal Provinces In fact Lawrence (1963 279) statedthat lsquoThis species resembles [Parapurcellia] silvicola fromZululand in the shape of the tarsus IVrsquo In particular it hasozophores oriented at 45 off the lateral margin rather thandorsal ozophores projections of the anterior gonostome walla small (~25mm) body length and no evidence of functionaleyes under light microscopy ndash unlike Purcellia which haslensless eyes situated in the bases of the ozophores Based onthese characters as well as several others discussed belowand consistent placement of Purcellia peregrinator withinParapurcellia in our phylogenetic analyses we reassignPurcellia peregrinator to Parapurcellia establishing the newcombination Parapurcellia peregrinator (Lawrence 1963)leaving the number of described species in Parapurcellia atfive Parapurcellia fissa (Lawrence 1939) and Parapurcelliasilvicola (Lawrence 1939) are distinctive due to their deeplylobed posterior tergites In the remaining two speciesParapurcellia monticola (Lawrence 1939) and Parapurcelliarumpiana (Lawrence 1933) the posterior tergites are relativelyunmodified and their morphologies are typical of small pettalidsalthough the former species is distinguishable from the otherSouth African species by its tall profile

The Natal Museum of South Africa (NMSA) kindlyprovided us with undescribed cyphophthalmid specimenscollected by several arachnologists between 1939 and 1985We identified two distinct morphospecies collected in theKnysna forest that straddles the border of the Eastern andWestern Cape Provinces which appear to belong in PurcelliaAn additional four morphospecies from localities in the EasternCape and KwazulundashNatal Provinces segregated years ago byW Starega who assigned tentative names to some of the speciesappear to belong in Parapurcellia We have chosen to use hisnames as much as possible to honour his early recognition of thedifferent morphologies These are described as new speciesbelow and our placement of them into their respective generais supported by morphological phylogenetic analysis

Speleosiro argasiformisPurcellia illustrans

Parapurcellia peregrinator

Parapurcellia fissa

Purcellia griswoldi sp nov

Parapurcellia amatola sp nov

Parapurcellia staregai sp nov

Parapurcellia minuta sp novParapurcellia natalia sp nov

Purcellia lawrencei sp nov

Parapurcellia monticolaParapurcellia rumpiana

Parapurcellia silvicola

Purcellia transvaalica

NORTHERN CAPE

WESTERN CAPE

EASTERN CAPE

FREE STATE

NORTH WEST

LIMPOPO

GAU-TENG

KWAZULU-NATAL

MPUMA-LANGA

NORTHERN CAPE

WESTERN CAPE

EASTERN CAPE

FREE STATE

NORTH WEST

LIMPOPO

GAU-TENG

KWAZULU-NATAL

MPUMA-LANGA

Parapurcellia convexa sp nov

Fig 1 Mapofpettalid type collection localities inSouthAfricaCircles indicate localities of speciesdescribedbeforethiswork Stars indicate new species described herein Square indicatesParapurcellia peregrinator (Lawrence 1963)new comb whose genus is reassigned herein

372 Invertebrate Systematics B L de Bivort and G Giribet

The NMSA collections along with most collections ofpreviously described species were made before the era ofmolecular phylogenetics Their long-term storage in 70ethanol has precluded the possibility of analysing the species-level relationships of the South African pettalids using molecular

sequences In order to investigate the phylogenetic relationshipsof the SouthAfricanmaterial we chose to perform aphylogeneticanalysis under the parsimony criterion using all availablemorphological data Attempts at lsquototal evidencersquo analysis haveproved successful in past analyses of the Cyphophthalmi

Table 1 Discrete character matrixCharacter codings are explained in Appendix 2

Taxa Characters000000000 1111111111 2222222222 3333333333 4444444444 5555555555 666

123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012

Parapurcellia staregai sp nov 032200011 031101100 011001100 00-000100 0112000100 1130100100 0

Parapurcellia amatolae sp nov 032200011 031100100 1101011000 00-0020100 0112000100 1130100100 0

Parapurcellia convexa sp nov 032200011 031100100 0101011100 00-0020100 0112000110 1120100101 0

Parapurcellia natalia sp nov 032200011 031000100 011101100 00-000100 0112000110 1130100100 0

Parapurcellia minuta sp nov 032200011 031101100 0101011000 00-0020110 0112000110 10-0100100 0

Parapurcellia peregrinator 032200011 021100100 111101110 00-00300 0112000110 1120100100 0

Parapurcellia silvicola 032200011 031100000 0101011100 00-0020100 01100110 0100100101 0

Parapurcellia monticola 032200011 021100000 0101011100 00-0020100 01100110 1130100100 0

Parapurcellia rumpiana 032200011 021001100 010111101 0-30100 01100110 110100100 0

Parapurcellia fissa 032200011 031100100 0101110 30100 0100110 130100100 0

Purcellia transvaalica 222200011 020101100 110101100 00-0020 0112100100 1140110100 0

Purcellia illustrans 222200011 000100000 1101011101 00-0020100 0112100100 0140110100 001

Purcellia griswoldi sp nov 222200010 020100100 1001011100 00-0020100 0112100100 1140110100 011

Purcellia lawrencei sp nov 222200011 030100100 110101100 00-0020100 0112100100 1140110100 0

Speleosiro argasiformis 022200011 000100000 1100011101 0-0020100 0111000100 0140110100 0

Rakaia isolata 232200011 000101000 110101100 00-0020000 0100110 0100101101 0

Rakaia lindsayi 232200011 000100000 1100010210 00-0020000 01100110 0100100101 0

Rakaia magna australis 232200011 000101000 1100011200 00-0020000 01100110 0100100100 0

Rakaia media 232200011 000100000 1101011200 00-0020000 011200110 0100100100 0

Rakaia solitaria 132200011 000100000 1101011200 00-0020000 01100110 0100101100 0

Rakaia sorenseni digitata 232200011 000100000 1101011100 00-0020050 010ndashndash00110 00-01-1000 0

Aoraki crypta 232200011 000100000 1110010101 00-0020000 011200110 01311-0000 0

Aoraki denticula 232200011 000100000 1110010201 00-0021000 011200110 01301-0001 0

Aoraki inerma 232200011 000100000 1110010201 00-0020000 011200110 01311-0000 0

Aoraki longitarsa 232200011 000100000 1110010201 00-0021000 0100110 0100100101 1

Aoraki tumidata 232200011 000101000 11001011 0-2 112100111 10-0101101 0

Karripurcellia harveyi 232200011 000100010 1110010210 00-0020000 010ndashndash10100 00-00-0000 0

Karripurcellia peckorum 232200011 000101010 1110010210 00-0020000 010ndashndash10100 00-00-0000 001

Karripurcellia sierwaldae 232200011 000101010 1100011200 00-0020000 010ndashndash10100 00-00-0000 0

Austropurcellia scoparia 232200011 000100000 1101011101 00-0030100 0114000110 01101-0000 11

Austropurcellia woodwardi 232200011 020100000 111101 00-0030100 011000110 0130100100 1

Chileogovea jocasta 132200011 001100000 1110010101 00-1140000 0112100100 00-00-0000 0

Chileogovea oedipus 132100011 001100000 1110010211 00-1141000 0112100101 10-00-0000 01

Pettalus lampetides 122200011 001100010 1110110001 00-0041001 010ndashndash02100 00-01-1000 01

Pettalus cimiciformis 122100011 00310001 1110010101 00-4100 010ndashndash02100 00-01-1000 0

Troglosiro aelleni 010210011 001000000 2110000201 0110030000 001003-00 00-00-0000 0

Troglosiro longifossa 010210011 0010100100 2110000101 0110030000 001003-00 00-00-0010 0

Troglosiro ninqua 010210011 0010100000 2110000000 0110030000 001003-00 00-00-0010 0

Huitaca ventralis 010200101 111-100010 2110010201 0110030001 001003-00 00-00-0000 00

Neogovea sp 010200111 1310101010 1101000 0 003-00 00-00-0000 0

Metagovea philipi 010200111 12110000 2110000101 0110030001 001003-00 00-00-0000 001

Cyphophthalmus duricorius 010211011 0020000101 0000010100 00-0010000 001103-01 10-00-0000 011

Parasiro coiffaiti 000201010 0010000101 0100010000 0111120000 000ndashndash01000 00-00-0000 010

Siro acaroides 010211010 0010000100 0000000000 00-1110000 0011103-00 10-01-0000 011

Siro rubens 010211011 0020000100 0000020100 00-0010000 0011103-01 00-00-0000 011

Suzukielus sauteri 012200011 001000100 0000020100 1101120100 0013000101 00-00-0000 011

Stylocellus ramblae 111200101 000010001 011011021 00-0151010 020ndashndash00000 00-00-0000 0

Stylocellus tambusisi 111200100 121010001 01100102 00-001010 020ndashndash00000 10-00-0000 0

Fangensis cavernarus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001

Fangensis spelaeus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001

Systematic revision of South African pettalids Invertebrate Systematics 373

Tab

le2

Con

tinu

ouscharactermatrixas

itap

pearsin

the11

weigh

tedcombinedan

alysis

Character

definitio

nsaregivenin

App

endix3

12

34

56

78

910

1112

1314

1516

1718

1920

2122

2324

2526

2728

2930

3132

3334

3536

3738

39

Parap

urcelliastaregaisp

nov

054

059

058

052

053

053

059

047

064

051

058

063

047

067

069

075

048

071

070

056

055

049

062

073

057

057

052

052

054

066

054

068

048

048

056

056

067

061

052

Parap

urcelliaam

atolaesp

nov

047

061

057

056

056

064

052

047

065

057

053

072

051

062

061

059

062

070

066

056

042

063

062

053

053

058

058

057

072

055

050

061

043

049

056

056

066

062

049

Parapurcelliaconvexa

spn

ov

048

069

058

075

067

066

049

053

060

051

050

078

081

063

074

051

064

062

071

059

056

065

057

059

045

066

085

075

071

070

068

052

057

051

056

056

061

060

054

Parapurcellianatalia

sp

nov

044

056

057

052

053

057

049

047

057

049

060

063

048

050

071

061

105

082

074

060

053

059

061

076

059

059

052

052

065

056

043

065

043

047

056

056

071

058

049

Parapurcelliaminuta

spn

ov

042

056

056

052

053

046

050

047

057

054

081

075

047

068

072

082

048

069

062

060

059

069

059

050

052

057

052

052

043

067

055

064

057

052

056

056

067

058

048

Parapurcelliaperegrinator

059

061

058

059

055

057

059

047

077

055

065

065

048

063

049

048

066

061

057

061

055

056

057

062

071

059

060

055

058

061

053

062

053

047

056

056

065

059

059

Parapurcelliasilvicola

055

063

063

076

064

049

062

050

062

062

060

072

077

053

079

050

065

079

070

059

060

065

064

064

057

071

080

067

047

097

059

068

040

059

056

056

064

058

070

Parapurcelliamonticola

054

061

059

061

053

056

045

053

073

052

058

074

045

055

064

073

050

070

059

057

065

062

058

064

061

063

063

052

058

054

066

049

074

051

056

056

066

063

062

Parap

urcelliarumpiana

051

059

056

057

053

050

040

053

067

078

046

073

075

057

073

080

037

074

071

057

061

058

054

056

059

059

060

052

049

062

066

039

061

046

056

056

075

060

055

Parapurcelliafissa

043

064

064

052

053

056

059

053

065

052

055

070

071

069

076

063

055

066

060

057

063

079

078

060

051

085

052

052

062

036

068

083

053

053

056

056

068

060

052

Purcelliatransvaalica

069

064

057

052

053

048

060

058

061

058

041

071

070

057

074

050

061

053

072

050

067

050

056

054

056

056

052

052

046

056

054

051

050

036

056

056

074

058

048

Purcelliaillustrans

072

063

055

052

053

052

060

087

081

056

061

063

062

061

077

049

059

075

071

051

065

058

051

078

087

054

052

052

050

089

096

053

076

036

056

056

078

061

042

Purcelliagriswoldisp

nov

062

063

057

057

053

043

062

055

067

053

065

066

062

044

064

068

050

057

058

048

075

059

055

053

058

058

057

052

041

061

061

060

060

040

056

056

075

057

045

Purcellialawrenceisp

nov

062

061

056

057

053

051

066

057

057

053

059

069

067

047

054

066

053

068

067

048

080

063

052

056

050

055

057

052

050

063

068

064

068

039

056

056

075

053

044

Speleosiro

argasiform

is097

077

060

075

059

073

091

048

071

051

049

044

063

039

056

055

068

031

035

045

108

046

054

047

050

057

064

056

061

053

056

060

057

047

056

056

070

058

043

Rakaiaisolate

059

077

067

102

058

069

066

061

059

063

067

077

077

070

062

046

077

054

068

052

062

062

072

061

043

076

104

058

071

066

061

068

059

062

056

056

058

064

099

Rakaialin

dsayi

061

063

062

083

053

059

053

057

054

066

071

056

071

077

067

086

046

079

059

051

057

071

058

057

061

070

082

052

061

063

037

055

037

056

056

056

062

066

062

Rakaiamag

naau

stralis

074

068

064

078

061

085

070

047

077

085

072

052

071

057

043

053

067

068

062

054

062

062

066

068

058

060

072

059

078

065

065

059

067

053

056

056

065

058

075

Rakaiamedia

062

070

062

063

057

055

077

062

066

052

066

058

070

081

066

087

041

065

064

049

057

057

068

058

068

061

062

057

054

058

054

077

057

064

056

056

061

063

077

Rakaiasolitaria

057

065

059

071

053

065

052

060

079

053

074

080

051

055

051

046

070

065

068

053

077

043

055

037

060

065

073

052

069

069

067

055

065

066

056

056

053

062

074

Rakaiasorensenidigitata

057

071

055

054

055

075

071

080

065

066

056

093

052

085

038

054

064

069

065

051

075

075

053

066

069

054

055

056

083

060

059

080

058

074

056

056

047

053

078

Aorakicrypta

071

079

061

058

059

085

069

088

041

059

049

059

043

072

059

047

071

060

064

052

060

037

051

039

053

071

057

058

081

067

062

061

053

039

056

056

077

075

086

Aorakidenticula

057

067

055

068

062

063

048

080

054

054

056

065

051

085

058

060

067

068

071

053

066

065

051

057

069

055

071

064

066

059

063

050

063

044

075

072

071

070

070

Aorakiinerma

067

066

057

063

063

062

056

070

075

056

044

066

065

077

053

051

066

060

059

054

073

066

053

053

063

057

062

063

060

054

060

050

059

062

056

056

059

072

074

Aorakilong

itarsa

066

076

073

098

109

065

058

069

056

058

083

056

049

062

028

049

061

063

064

050

061

059

066

079

060

091

094

107

064

076

062

055

058

064

056

056

059

129

062

Aorakitumidata

064

072

058

081

069

079

070

072

063

068

071

073

079

083

054

080

060

073

070

050

062

072

055

054

062

060

083

070

081

085

062

070

050

070

075

096

080

059

071

Karripurcellia

harveyi

067

052

058

061

066

051

079

089

062

052

069

056

073

060

074

075

037

050

043

055

056

063

060

069

091

052

060

065

049

066

050

072

048

067

056

056

047

057

058

Karripurcellia

peckorum

068

062

057

071

068

069

062

070

051

094

057

064

070

057

052

045

071

046

037

055

060

064

057

059

076

054

069

067

064

070

046

053

037

068

056

056

047

056

056

Karripurcellia

sierwaldae

056

056

057

052

053

042

062

061

067

088

056

061

076

063

070

057

060

043

032

055

062

069

062

067

068

053

052

052

039

067

057

064

051

073

064

087

047

057

060

Austropurcelliascoparia

055

050

054

052

053

067

052

068

054

051

045

065

050

064

057

044

071

065

074

059

071

072

051

054

084

053

052

052

071

052

082

055

088

062

056

056

061

091

053

Austrop

urcelliawoo

dwardi

054

065

058

052

058

056

061

069

053

053

042

066

049

066

068

053

057

062

068

054

063

047

049

036

050

067

052

060

056

065

079

064

068

059

056

056

047

061

053

Chileogovea

jocasta

049

053

055

052

054

058

050

057

055

049

047

049

058

060

055

063

050

050

058

051

074

055

053

053

060

053

052

055

061

057

069

057

068

066

082

088

082

053

069

Chileog

ovea

Oedipus

070

053

054

052

053

064

070

074

057

062

053

052

043

064

054

051

058

072

078

053

071

012

049

043

054

052

052

052

061

052

063

061

070

062

092

091

083

051

048

Pettaluslampetid

es063

066

078

052

092

067

057

058

053

055

063

039

048

055

057

063

064

068

068

046

064

050

095

090

044

084

052

093

065

042

050

052

066

074

056

056

047

052

055

Pettaluscimicifo

rmis

100

110

140

066

095

089

082

076

066

058

074

033

039

021

057

058

058

056

046

041

056

085

125

064

044

116

059

074

070

039

047

055

071

068

056

056

047

050

061

Troglosiroaelleni

059

053

059

054

066

066

064

057

054

053

052

040

067

066

072

060

064

044

053

076

045

064

066

075

062

054

054

067

068

054

061

066

064

074

056

056

047

055

058

Troglosirolong

ifossa

051

045

055

052

073

064

048

070

049

060

057

042

057

053

069

043

069

053

064

079

041

059

055

061

066

052

052

081

070

049

049

052

055

074

056

056

047

056

053

Troglosironinq

ua059

045

057

053

086

068

049

074

049

066

048

044

074

054

064

052

063

048

059

076

044

054

063

072

068

052

054

092

069

062

076

048

070

074

056

056

047

057

072

Huitaca

ventralis

075

045

057

052

060

036

084

047

105

094

063

054

058

067

072

068

054

040

056

075

057

046

058

070

053

052

052

058

039

045

038

057

047

074

056

056

047

052

048

Neogo

veasp

067

056

058

065

070

031

070

075

081

099

042

055

067

047

063

068

042

031

045

074

050

038

061

073

061

052

063

069

033

035

035

055

044

074

056

056

047

050

045

Metagovea

philipi

045

045

054

052

053

041

045

057

065

055

056

056

066

051

053

054

056

048

062

078

057

057

053

050

031

052

052

052

036

042

056

048

073

074

056

056

047

053

068

Cypho

phthalmus

duricorius

050

045

055

052

053

058

041

050

039

060

067

044

044

042

027

050

072

058

065

078

039

065

057

071

065

052

052

052

062

059

067

044

072

074

056

056

047

056

065

Parasirocoiffaiti

044

045

055

054

055

058

056

047

050

052

079

050

041

060

056

080

051

064

061

086

042

064

060

062

058

052

056

057

068

053

052

081

063

068

056

056

055

053

064

Siro

acaroids

046

055

054

052

053

055

053

049

044

055

047

051

067

060

070

051

057

058

071

069

048

068

053

044

061

053

052

052

058

062

061

062

055

051

056

056

071

055

068

Siro

rubens

049

045

056

052

053

042

077

050

041

055

065

039

057

053

062

082

037

047

052

072

035

073

059

052

086

052

052

052

036

054

085

102

100

066

119

102

056

050

049

Suzukielus

sauteri

059

045

056

052

053

060

062

048

050

051

063

068

069

065

064

066

056

082

071

075

048

071

056

067

057

052

052

052

062

079

072

066

063

071

098

091

047

056

061

Stylocellusramblae

052

045

055

052

053

064

050

060

052

048

083

058

047

056

053

055

076

045

046

092

068

060

054

046

048

052

052

052

067

053

058

053

074

074

056

056

047

053

060

Stylocellustambusisi

074

045

057

052

053

066

049

059

046

067

058

052

050

041

029

059

076

047

056

082

066

060

057

070

056

052

052

052

061

047

055

041

065

074

056

056

047

052

062

Fan

gensiscavernarus

081

064

065

052

053

073

046

047

049

065

080

054

076

053

065

048

079

048

026

075

061

073

065

078

081

059

052

052

066

063

049

038

051

074

056

056

047

053

048

Fan

gensisspelaeus

073

045

057

052

053

078

093

047

050

061

093

065

077

056

064

076

053

068

030

076

054

069

056

069

066

052

052

052

076

068

055

086

061

074

056

056

047

057

054

374 Invertebrate Systematics B L de Bivort and G Giribet

4041

4243

4445

4647

4849

5051

5253

5455

5657

5859

6061

6263

6465

6667

6869

7071

7273

7475

7677

78

Parap

urcelliastaregaisp

nov

054

049

048

053

061

054

046

048

065

057

060

056

057

059

069

055

074

063

048

049

055

058

069

062

068

080

057

066

053

057

056

052

068

059

066

054

070

051

062

Parap

urcelliaam

atolaesp

nov

047

058

055

044

066

054

046

050

081

058

068

069

065

064

068

055

068

055

049

032

047

064

064

070

066

059

053

073

046

043

035

075

066

053

068

048

065

062

056

Parapurcelliaconvexa

spn

ov

036

030

068

095

053

054

055

050

064

057

053

052

056

065

070

055

069

066

055

062

053

069

052

075

070

059

063

067

049

055

069

062

077

034

062

057

071

052

062

Parapurcellianatalia

sp

nov

050

046

050

051

054

054

046

046

079

060

056

066

072

063

064

055

062

060

057

058

046

068

053

063

063

069

042

063

045

054

073

050

062

073

056

054

060

048

054

Parap

urcelliaminuta

spn

ov

046

046

054

041

030

054

047

049

075

060

059

050

058

059

066

055

062

062

057

040

062

066

072

061

062

076

044

064

052

056

058

076

095

069

053

058

062

057

054

Parap

urcelliaperegrinator

058

058

058

050

049

054

046

054

071

063

049

056

070

062

060

055

062

060

057

070

053

063

073

077

079

056

052

078

049

061

058

057

064

079

052

048

066

055

055

Parapurcelliasilvicola

052

054

062

050

044

054

050

048

064

058

063

059

055

062

069

055

064

061

052

051

062

067

072

070

066

019

045

064

059

059

048

061

074

066

065

073

070

064

057

Parapurcelliamonticola

067

073

086

060

055

054

053

051

066

057

050

062

071

063

070

055

060

065

058

053

044

064

061

039

072

075

047

080

048

061

056

047

066

061

066

047

061

053

056

Parap

urcelliarumpian

a040

035

072

081

058

054

055

045

075

057

048

067

074

063

065

055

082

059

056

072

052

066

051

064

073

068

049

076

073

066

056

058

070

035

054

090

078

093

058

Parapurcelliafissa

060

057

065

048

059

054

057

060

063

057

049

056

066

063

060

055

062

061

051

064

056

065

062

038

074

051

057

079

050

059

043

048

069

061

067

051

071

055

060

Purcelliatransvaalica

064

068

060

051

053

054

056

057

062

055

044

054

065

055

069

055

077

068

052

054

044

068

069

066

064

058

060

058

060

071

050

043

054

051

073

055

054

056

058

Purcelliaillustrans

111

080

101

074

037

054

082

067

066

052

047

046

060

066

057

055

056

056

055

047

052

066

063

057

066

048

055

077

052

068

051

043

059

048

053

047

072

054

059

Purcelliagriswoldisp

nov

067

078

063

048

049

054

054

049

065

057

045

050

066

061

065

055

053

061

059

046

050

063

074

056

070

071

070

066

058

070

035

050

051

069

049

058

077

052

056

Purcellialawrenceisp

nov

071

081

067

052

046

054

056

052

089

054

046

040

054

061

066

055

059

059

069

047

038

074

067

054

072

068

056

056

056

065

051

050

050

050

042

062

068

052

057

Speleosiro

argasiform

is057

056

054

045

060

054

047

044

055

066

041

058

078

061

045

055

058

061

057

044

076

049

090

079

055

054

085

058

077

089

050

052

051

070

043

041

050

048

060

Rakaiaisolate

066

075

061

060

062

054

061

079

032

073

059

069

071

063

059

055

066

060

066

044

061

052

039

053

069

056

038

058

049

050

049

077

071

058

077

062

065

064

059

Rakaialin

dsayi

050

060

058

056

068

054

057

054

070

057

067

065

060

067

060

055

055

056

068

059

073

059

060

044

078

074

049

053

060

059

050

064

057

085

059

055

064

066

058

Rakaiamag

naau

stralis

060

047

055

055

055

054

046

052

059

078

064

074

072

066

054

055

061

064

073

060

060

064

050

056

070

047

058

068

062

052

053

074

049

049

058

061

070

077

058

Rakaiamedia

060

082

041

052

062

054

061

052

065

078

064

082

075

061

058

055

060

062

075

060

047

059

043

059

063

071

060

064

049

037

076

056

070

050

061

063

070

051

060

Rakaiasolitaria

065

071

056

052

051

054

060

066

068

067

074

075

063

069

053

055

078

056

065

060

053

064

047

055

063

051

057

081

050

048

074

072

056

065

075

069

073

052

058

Rakaiasorensenidigitata

076

071

066

060

034

054

087

052

064

060

049

030

039

084

055

055

065

065

065

060

068

057

072

072

072

051

067

069

061

062

061

078

076

078

077

073

071

070

048

Aorakicrypta

071

069

053

060

054

054

082

048

068

067

068

063

051

069

055

055

059

073

071

054

064

047

075

078

071

056

080

040

068

049

067

081

042

064

068

041

047

057

048

Aorakidenticula

075

068

054

061

068

054

085

055

052

057

068

055

048

077

054

055

070

060

056

065

047

073

071

049

074

066

053

055

057

035

085

061

070

067

058

040

059

054

060

Aorakiinerma

069

070

071

078

065

054

069

046

068

060

064

063

052

072

050

055

081

064

054

050

058

060

062

072

042

055

057

073

070

068

054

052

047

043

079

046

057

055

061

Aorakilong

itarsa

069

066

057

062

073

054

067

078

064

059

075

063

056

071

056

055

065

073

057

069

073

043

074

071

064

050

071

054

064

049

071

078

050

052

062

056

063

057

062

Aorakitumidata

066

062

067

047

063

054

073

062

066

055

074

087

071

070

065

055

051

065

072

062

095

023

074

082

079

081

077

063

082

060

064

105

068

085

055

059

071

069

069

Karripurcellia

harveyi

050

061

072

074

086

054

084

076

057

054

047

047

064

062

057

055

056

054

061

077

075

053

060

073

053

047

066

059

064

070

054

051

055

061

045

052

054

050

060

Karripurcellia

peckorum

046

050

066

077

087

054

067

056

050

067

045

047

062

064

056

055

059

059

074

071

069

056

061

059

063

045

055

049

082

067

054

055

042

054

081

071

051

090

055

Karripurcellia

sierwaldae

050

052

064

067

068

054

062

061

060

050

060

044

044

064

065

055

052

060

051

060

060

058

057

064

061

060

063

070

079

074

051

040

045

058

056

081

057

097

048

Austropurcelliascoparia

079

078

069

066

053

054

074

064

051

052

080

056

037

089

038

055

056

057

080

050

057

076

067

062

060

048

053

056

052

062

064

060

061

068

062

066

062

052

063

Austrop

urcelliawoo

dwardi

072

067

074

061

054

054

075

079

049

062

075

067

047

074

036

055

058

058

053

054

051

086

061

052

056

059

038

054

054

065

060

046

058

049

063

080

067

054

048

Chileogovea

jocasta

067

064

065

054

054

054

061

052

054

055

069

066

052

069

045

055

049

058

057

057

063

066

042

053

059

068

068

059

056

074

054

056

057

060

043

057

046

050

048

Chileog

ovea

oedipu

s064

068

070

070

048

054

070

076

081

056

070

056

046

069

056

055

054

059

054

085

072

060

047

044

076

057

066

055

072

074

048

064

054

076

046

052

041

060

048

Pettaluslampetid

es063

055

059

058

058

054

057

051

063

063

053

077

084

052

075

055

058

051

057

058

054

052

051

057

060

063

062

052

073

067

054

051

039

075

083

040

034

053

061

Pettaluscimicifo

rmis

063

061

050

062

048

062

060

058

044

065

044

054

079

048

070

055

042

048

069

049

060

051

057

051

058

062

082

054

051

061

077

059

060

053

072

046

062

051

057

Troglosiroaelleni

059

056

029

089

066

085

058

054

066

046

078

071

050

062

045

090

056

060

068

076

055

055

066

058

045

064

076

054

064

065

059

049

055

055

046

049

037

052

060

Troglosirolongifo

ssa

059

055

058

070

077

085

077

053

061

045

069

071

058

048

054

093

038

065

070

085

059

056

052

082

044

051

071

049

056

045

075

053

050

053

038

063

073

064

048

Troglosironinq

ua066

051

072

073

079

083

077

061

060

041

068

067

053

045

064

092

062

059

058

067

059

053

070

075

052

061

082

049

075

076

046

055

048

051

036

053

045

068

048

Huitaca

ventralis

045

045

040

050

059

085

046

058

056

041

049

070

083

032

075

081

031

044

049

051

062

061

070

030

046

062

071

090

108

096

085

062

043

073

043

063

033

084

048

Neogo

veasp

042

067

051

086

073

087

071

066

031

046

039

066

087

037

059

055

066

130

104

061

066

071

062

039

042

072

075

076

074

077

079

055

050

036

084

064

038

095

061

Metag

ovea

philipi

057

068

061

073

063

085

062

044

040

041

066

069

059

039

057

055

061

044

080

064

056

085

058

045

035

059

041

075

059

063

053

049

046

047

065

054

034

059

048

Cypho

phthalmus

duricorius

060

046

053

056

061

086

050

058

065

056

053

064

073

054

051

087

060

064

057

082

060

061

060

065

048

047

064

036

055

052

070

059

066

065

071

077

072

064

078

Parasirocoiffaiti

056

064

071

050

066

081

046

074

063

060

089

070

051

051

055

078

055

065

053

062

063

060

032

073

042

079

053

052

044

048

078

068

088

061

060

077

070

055

048

Siro

acaroides

052

040

038

053

081

085

049

077

058

060

057

079

073

055

059

096

032

061

049

068

045

071

040

057

055

059

058

043

065

064

063

065

066

050

059

067

056

059

048

Siro

rubens

063

050

064

053

063

085

050

069

056

049

076

069

054

060

051

087

073

067

078

089

056

064

034

060

046

084

064

038

056

054

068

064

076

049

045

063

068

058

073

Suzukielus

sauteri

064

047

063

048

073

054

047

101

069

053

063

050

048

052

075

055

060

062

067

077

059

058

042

069

068

069

052

050

049

048

075

061

066

056

069

065

074

050

068

Stylocellusramblae

060

072

062

053

058

054

062

073

041

098

074

042

040

035

071

055

025

044

038

053

105

053

072

051

030

061

082

052

056

052

067

075

059

072

063

066

048

047

099

Stylocellustambusisi

056

063

061

052

060

054

056

069

047

096

073

050

041

040

067

055

063

044

038

042

064

060

074

050

036

048

063

044

060

049

067

060

058

078

057

085

046

062

087

Fan

gensiscavernarus

047

044

040

056

075

054

046

070

036

085

050

042

058

041

084

055

077

044

038

066

076

035

052

064

049

055

070

046

051

044

056

067

062

071

050

058

060

057

096

Fan

gensisspelaeus

049

061

044

055

079

054

046

070

047

086

046

035

058

042

105

055

087

044

038

065

077

029

068

065

060

084

050

047

051

045

078

070

083

057

054

071

065

056

096

Systematic revision of South African pettalids Invertebrate Systematics 375

combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)

Methods and abbreviations

Specimens used in this study were provided by the followinginstitutions

AMNH American Museum of Natural History New York(USA)

BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard

University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand

Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo

Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg

(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg

Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)

Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg

(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen

Copenhagen (Denmark)

Specimen handling and illustration

In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was

photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy

Character coding

Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3

Phylogenetic analyses

The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)

The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the

376 Invertebrate Systematics B L de Bivort and G Giribet

continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data

Taxonomy

Order OPILIONES Sundevall

Suborder CYPHOPHTHALMI Simon

Family PETTALIDAE Simon

Genus Speleosiro Lawrence 1931

Diagnosis

Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII

Type species

Speleosiro argasiformis Lawrence 1931

Species included

Speleosiro argasiformis Lawrence 1931

Distribution

SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province

Remarks

This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement

Genus Purcellia Hansen amp Soslashrensen 1904

Diagnosis

Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)

Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)

Type species

Purcellia illustrans Hansen amp Soslashrensen 1904

Species included

Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov

Distribution

South Africa Limpopo Eastern Cape and Western CapeProvinces

Remarks

This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa

Purcellia griswoldi sp nov

(Figs 2 3 Table 3)

Material examined

Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985

Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981

Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981

Systematic revision of South African pettalids Invertebrate Systematics 377

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 3: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

The NMSA collections along with most collections ofpreviously described species were made before the era ofmolecular phylogenetics Their long-term storage in 70ethanol has precluded the possibility of analysing the species-level relationships of the South African pettalids using molecular

sequences In order to investigate the phylogenetic relationshipsof the SouthAfricanmaterial we chose to perform aphylogeneticanalysis under the parsimony criterion using all availablemorphological data Attempts at lsquototal evidencersquo analysis haveproved successful in past analyses of the Cyphophthalmi

Table 1 Discrete character matrixCharacter codings are explained in Appendix 2

Taxa Characters000000000 1111111111 2222222222 3333333333 4444444444 5555555555 666

123456789 0123456789 0123456789 0123456789 0123456789 0123456789 012

Parapurcellia staregai sp nov 032200011 031101100 011001100 00-000100 0112000100 1130100100 0

Parapurcellia amatolae sp nov 032200011 031100100 1101011000 00-0020100 0112000100 1130100100 0

Parapurcellia convexa sp nov 032200011 031100100 0101011100 00-0020100 0112000110 1120100101 0

Parapurcellia natalia sp nov 032200011 031000100 011101100 00-000100 0112000110 1130100100 0

Parapurcellia minuta sp nov 032200011 031101100 0101011000 00-0020110 0112000110 10-0100100 0

Parapurcellia peregrinator 032200011 021100100 111101110 00-00300 0112000110 1120100100 0

Parapurcellia silvicola 032200011 031100000 0101011100 00-0020100 01100110 0100100101 0

Parapurcellia monticola 032200011 021100000 0101011100 00-0020100 01100110 1130100100 0

Parapurcellia rumpiana 032200011 021001100 010111101 0-30100 01100110 110100100 0

Parapurcellia fissa 032200011 031100100 0101110 30100 0100110 130100100 0

Purcellia transvaalica 222200011 020101100 110101100 00-0020 0112100100 1140110100 0

Purcellia illustrans 222200011 000100000 1101011101 00-0020100 0112100100 0140110100 001

Purcellia griswoldi sp nov 222200010 020100100 1001011100 00-0020100 0112100100 1140110100 011

Purcellia lawrencei sp nov 222200011 030100100 110101100 00-0020100 0112100100 1140110100 0

Speleosiro argasiformis 022200011 000100000 1100011101 0-0020100 0111000100 0140110100 0

Rakaia isolata 232200011 000101000 110101100 00-0020000 0100110 0100101101 0

Rakaia lindsayi 232200011 000100000 1100010210 00-0020000 01100110 0100100101 0

Rakaia magna australis 232200011 000101000 1100011200 00-0020000 01100110 0100100100 0

Rakaia media 232200011 000100000 1101011200 00-0020000 011200110 0100100100 0

Rakaia solitaria 132200011 000100000 1101011200 00-0020000 01100110 0100101100 0

Rakaia sorenseni digitata 232200011 000100000 1101011100 00-0020050 010ndashndash00110 00-01-1000 0

Aoraki crypta 232200011 000100000 1110010101 00-0020000 011200110 01311-0000 0

Aoraki denticula 232200011 000100000 1110010201 00-0021000 011200110 01301-0001 0

Aoraki inerma 232200011 000100000 1110010201 00-0020000 011200110 01311-0000 0

Aoraki longitarsa 232200011 000100000 1110010201 00-0021000 0100110 0100100101 1

Aoraki tumidata 232200011 000101000 11001011 0-2 112100111 10-0101101 0

Karripurcellia harveyi 232200011 000100010 1110010210 00-0020000 010ndashndash10100 00-00-0000 0

Karripurcellia peckorum 232200011 000101010 1110010210 00-0020000 010ndashndash10100 00-00-0000 001

Karripurcellia sierwaldae 232200011 000101010 1100011200 00-0020000 010ndashndash10100 00-00-0000 0

Austropurcellia scoparia 232200011 000100000 1101011101 00-0030100 0114000110 01101-0000 11

Austropurcellia woodwardi 232200011 020100000 111101 00-0030100 011000110 0130100100 1

Chileogovea jocasta 132200011 001100000 1110010101 00-1140000 0112100100 00-00-0000 0

Chileogovea oedipus 132100011 001100000 1110010211 00-1141000 0112100101 10-00-0000 01

Pettalus lampetides 122200011 001100010 1110110001 00-0041001 010ndashndash02100 00-01-1000 01

Pettalus cimiciformis 122100011 00310001 1110010101 00-4100 010ndashndash02100 00-01-1000 0

Troglosiro aelleni 010210011 001000000 2110000201 0110030000 001003-00 00-00-0000 0

Troglosiro longifossa 010210011 0010100100 2110000101 0110030000 001003-00 00-00-0010 0

Troglosiro ninqua 010210011 0010100000 2110000000 0110030000 001003-00 00-00-0010 0

Huitaca ventralis 010200101 111-100010 2110010201 0110030001 001003-00 00-00-0000 00

Neogovea sp 010200111 1310101010 1101000 0 003-00 00-00-0000 0

Metagovea philipi 010200111 12110000 2110000101 0110030001 001003-00 00-00-0000 001

Cyphophthalmus duricorius 010211011 0020000101 0000010100 00-0010000 001103-01 10-00-0000 011

Parasiro coiffaiti 000201010 0010000101 0100010000 0111120000 000ndashndash01000 00-00-0000 010

Siro acaroides 010211010 0010000100 0000000000 00-1110000 0011103-00 10-01-0000 011

Siro rubens 010211011 0020000100 0000020100 00-0010000 0011103-01 00-00-0000 011

Suzukielus sauteri 012200011 001000100 0000020100 1101120100 0013000101 00-00-0000 011

Stylocellus ramblae 111200101 000010001 011011021 00-0151010 020ndashndash00000 00-00-0000 0

Stylocellus tambusisi 111200100 121010001 01100102 00-001010 020ndashndash00000 10-00-0000 0

Fangensis cavernarus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001

Fangensis spelaeus 011200100 001010101 0100010210 00-0051010 121100000 00-00-0000 001

Systematic revision of South African pettalids Invertebrate Systematics 373

Tab

le2

Con

tinu

ouscharactermatrixas

itap

pearsin

the11

weigh

tedcombinedan

alysis

Character

definitio

nsaregivenin

App

endix3

12

34

56

78

910

1112

1314

1516

1718

1920

2122

2324

2526

2728

2930

3132

3334

3536

3738

39

Parap

urcelliastaregaisp

nov

054

059

058

052

053

053

059

047

064

051

058

063

047

067

069

075

048

071

070

056

055

049

062

073

057

057

052

052

054

066

054

068

048

048

056

056

067

061

052

Parap

urcelliaam

atolaesp

nov

047

061

057

056

056

064

052

047

065

057

053

072

051

062

061

059

062

070

066

056

042

063

062

053

053

058

058

057

072

055

050

061

043

049

056

056

066

062

049

Parapurcelliaconvexa

spn

ov

048

069

058

075

067

066

049

053

060

051

050

078

081

063

074

051

064

062

071

059

056

065

057

059

045

066

085

075

071

070

068

052

057

051

056

056

061

060

054

Parapurcellianatalia

sp

nov

044

056

057

052

053

057

049

047

057

049

060

063

048

050

071

061

105

082

074

060

053

059

061

076

059

059

052

052

065

056

043

065

043

047

056

056

071

058

049

Parapurcelliaminuta

spn

ov

042

056

056

052

053

046

050

047

057

054

081

075

047

068

072

082

048

069

062

060

059

069

059

050

052

057

052

052

043

067

055

064

057

052

056

056

067

058

048

Parapurcelliaperegrinator

059

061

058

059

055

057

059

047

077

055

065

065

048

063

049

048

066

061

057

061

055

056

057

062

071

059

060

055

058

061

053

062

053

047

056

056

065

059

059

Parapurcelliasilvicola

055

063

063

076

064

049

062

050

062

062

060

072

077

053

079

050

065

079

070

059

060

065

064

064

057

071

080

067

047

097

059

068

040

059

056

056

064

058

070

Parapurcelliamonticola

054

061

059

061

053

056

045

053

073

052

058

074

045

055

064

073

050

070

059

057

065

062

058

064

061

063

063

052

058

054

066

049

074

051

056

056

066

063

062

Parap

urcelliarumpiana

051

059

056

057

053

050

040

053

067

078

046

073

075

057

073

080

037

074

071

057

061

058

054

056

059

059

060

052

049

062

066

039

061

046

056

056

075

060

055

Parapurcelliafissa

043

064

064

052

053

056

059

053

065

052

055

070

071

069

076

063

055

066

060

057

063

079

078

060

051

085

052

052

062

036

068

083

053

053

056

056

068

060

052

Purcelliatransvaalica

069

064

057

052

053

048

060

058

061

058

041

071

070

057

074

050

061

053

072

050

067

050

056

054

056

056

052

052

046

056

054

051

050

036

056

056

074

058

048

Purcelliaillustrans

072

063

055

052

053

052

060

087

081

056

061

063

062

061

077

049

059

075

071

051

065

058

051

078

087

054

052

052

050

089

096

053

076

036

056

056

078

061

042

Purcelliagriswoldisp

nov

062

063

057

057

053

043

062

055

067

053

065

066

062

044

064

068

050

057

058

048

075

059

055

053

058

058

057

052

041

061

061

060

060

040

056

056

075

057

045

Purcellialawrenceisp

nov

062

061

056

057

053

051

066

057

057

053

059

069

067

047

054

066

053

068

067

048

080

063

052

056

050

055

057

052

050

063

068

064

068

039

056

056

075

053

044

Speleosiro

argasiform

is097

077

060

075

059

073

091

048

071

051

049

044

063

039

056

055

068

031

035

045

108

046

054

047

050

057

064

056

061

053

056

060

057

047

056

056

070

058

043

Rakaiaisolate

059

077

067

102

058

069

066

061

059

063

067

077

077

070

062

046

077

054

068

052

062

062

072

061

043

076

104

058

071

066

061

068

059

062

056

056

058

064

099

Rakaialin

dsayi

061

063

062

083

053

059

053

057

054

066

071

056

071

077

067

086

046

079

059

051

057

071

058

057

061

070

082

052

061

063

037

055

037

056

056

056

062

066

062

Rakaiamag

naau

stralis

074

068

064

078

061

085

070

047

077

085

072

052

071

057

043

053

067

068

062

054

062

062

066

068

058

060

072

059

078

065

065

059

067

053

056

056

065

058

075

Rakaiamedia

062

070

062

063

057

055

077

062

066

052

066

058

070

081

066

087

041

065

064

049

057

057

068

058

068

061

062

057

054

058

054

077

057

064

056

056

061

063

077

Rakaiasolitaria

057

065

059

071

053

065

052

060

079

053

074

080

051

055

051

046

070

065

068

053

077

043

055

037

060

065

073

052

069

069

067

055

065

066

056

056

053

062

074

Rakaiasorensenidigitata

057

071

055

054

055

075

071

080

065

066

056

093

052

085

038

054

064

069

065

051

075

075

053

066

069

054

055

056

083

060

059

080

058

074

056

056

047

053

078

Aorakicrypta

071

079

061

058

059

085

069

088

041

059

049

059

043

072

059

047

071

060

064

052

060

037

051

039

053

071

057

058

081

067

062

061

053

039

056

056

077

075

086

Aorakidenticula

057

067

055

068

062

063

048

080

054

054

056

065

051

085

058

060

067

068

071

053

066

065

051

057

069

055

071

064

066

059

063

050

063

044

075

072

071

070

070

Aorakiinerma

067

066

057

063

063

062

056

070

075

056

044

066

065

077

053

051

066

060

059

054

073

066

053

053

063

057

062

063

060

054

060

050

059

062

056

056

059

072

074

Aorakilong

itarsa

066

076

073

098

109

065

058

069

056

058

083

056

049

062

028

049

061

063

064

050

061

059

066

079

060

091

094

107

064

076

062

055

058

064

056

056

059

129

062

Aorakitumidata

064

072

058

081

069

079

070

072

063

068

071

073

079

083

054

080

060

073

070

050

062

072

055

054

062

060

083

070

081

085

062

070

050

070

075

096

080

059

071

Karripurcellia

harveyi

067

052

058

061

066

051

079

089

062

052

069

056

073

060

074

075

037

050

043

055

056

063

060

069

091

052

060

065

049

066

050

072

048

067

056

056

047

057

058

Karripurcellia

peckorum

068

062

057

071

068

069

062

070

051

094

057

064

070

057

052

045

071

046

037

055

060

064

057

059

076

054

069

067

064

070

046

053

037

068

056

056

047

056

056

Karripurcellia

sierwaldae

056

056

057

052

053

042

062

061

067

088

056

061

076

063

070

057

060

043

032

055

062

069

062

067

068

053

052

052

039

067

057

064

051

073

064

087

047

057

060

Austropurcelliascoparia

055

050

054

052

053

067

052

068

054

051

045

065

050

064

057

044

071

065

074

059

071

072

051

054

084

053

052

052

071

052

082

055

088

062

056

056

061

091

053

Austrop

urcelliawoo

dwardi

054

065

058

052

058

056

061

069

053

053

042

066

049

066

068

053

057

062

068

054

063

047

049

036

050

067

052

060

056

065

079

064

068

059

056

056

047

061

053

Chileogovea

jocasta

049

053

055

052

054

058

050

057

055

049

047

049

058

060

055

063

050

050

058

051

074

055

053

053

060

053

052

055

061

057

069

057

068

066

082

088

082

053

069

Chileog

ovea

Oedipus

070

053

054

052

053

064

070

074

057

062

053

052

043

064

054

051

058

072

078

053

071

012

049

043

054

052

052

052

061

052

063

061

070

062

092

091

083

051

048

Pettaluslampetid

es063

066

078

052

092

067

057

058

053

055

063

039

048

055

057

063

064

068

068

046

064

050

095

090

044

084

052

093

065

042

050

052

066

074

056

056

047

052

055

Pettaluscimicifo

rmis

100

110

140

066

095

089

082

076

066

058

074

033

039

021

057

058

058

056

046

041

056

085

125

064

044

116

059

074

070

039

047

055

071

068

056

056

047

050

061

Troglosiroaelleni

059

053

059

054

066

066

064

057

054

053

052

040

067

066

072

060

064

044

053

076

045

064

066

075

062

054

054

067

068

054

061

066

064

074

056

056

047

055

058

Troglosirolong

ifossa

051

045

055

052

073

064

048

070

049

060

057

042

057

053

069

043

069

053

064

079

041

059

055

061

066

052

052

081

070

049

049

052

055

074

056

056

047

056

053

Troglosironinq

ua059

045

057

053

086

068

049

074

049

066

048

044

074

054

064

052

063

048

059

076

044

054

063

072

068

052

054

092

069

062

076

048

070

074

056

056

047

057

072

Huitaca

ventralis

075

045

057

052

060

036

084

047

105

094

063

054

058

067

072

068

054

040

056

075

057

046

058

070

053

052

052

058

039

045

038

057

047

074

056

056

047

052

048

Neogo

veasp

067

056

058

065

070

031

070

075

081

099

042

055

067

047

063

068

042

031

045

074

050

038

061

073

061

052

063

069

033

035

035

055

044

074

056

056

047

050

045

Metagovea

philipi

045

045

054

052

053

041

045

057

065

055

056

056

066

051

053

054

056

048

062

078

057

057

053

050

031

052

052

052

036

042

056

048

073

074

056

056

047

053

068

Cypho

phthalmus

duricorius

050

045

055

052

053

058

041

050

039

060

067

044

044

042

027

050

072

058

065

078

039

065

057

071

065

052

052

052

062

059

067

044

072

074

056

056

047

056

065

Parasirocoiffaiti

044

045

055

054

055

058

056

047

050

052

079

050

041

060

056

080

051

064

061

086

042

064

060

062

058

052

056

057

068

053

052

081

063

068

056

056

055

053

064

Siro

acaroids

046

055

054

052

053

055

053

049

044

055

047

051

067

060

070

051

057

058

071

069

048

068

053

044

061

053

052

052

058

062

061

062

055

051

056

056

071

055

068

Siro

rubens

049

045

056

052

053

042

077

050

041

055

065

039

057

053

062

082

037

047

052

072

035

073

059

052

086

052

052

052

036

054

085

102

100

066

119

102

056

050

049

Suzukielus

sauteri

059

045

056

052

053

060

062

048

050

051

063

068

069

065

064

066

056

082

071

075

048

071

056

067

057

052

052

052

062

079

072

066

063

071

098

091

047

056

061

Stylocellusramblae

052

045

055

052

053

064

050

060

052

048

083

058

047

056

053

055

076

045

046

092

068

060

054

046

048

052

052

052

067

053

058

053

074

074

056

056

047

053

060

Stylocellustambusisi

074

045

057

052

053

066

049

059

046

067

058

052

050

041

029

059

076

047

056

082

066

060

057

070

056

052

052

052

061

047

055

041

065

074

056

056

047

052

062

Fan

gensiscavernarus

081

064

065

052

053

073

046

047

049

065

080

054

076

053

065

048

079

048

026

075

061

073

065

078

081

059

052

052

066

063

049

038

051

074

056

056

047

053

048

Fan

gensisspelaeus

073

045

057

052

053

078

093

047

050

061

093

065

077

056

064

076

053

068

030

076

054

069

056

069

066

052

052

052

076

068

055

086

061

074

056

056

047

057

054

374 Invertebrate Systematics B L de Bivort and G Giribet

4041

4243

4445

4647

4849

5051

5253

5455

5657

5859

6061

6263

6465

6667

6869

7071

7273

7475

7677

78

Parap

urcelliastaregaisp

nov

054

049

048

053

061

054

046

048

065

057

060

056

057

059

069

055

074

063

048

049

055

058

069

062

068

080

057

066

053

057

056

052

068

059

066

054

070

051

062

Parap

urcelliaam

atolaesp

nov

047

058

055

044

066

054

046

050

081

058

068

069

065

064

068

055

068

055

049

032

047

064

064

070

066

059

053

073

046

043

035

075

066

053

068

048

065

062

056

Parapurcelliaconvexa

spn

ov

036

030

068

095

053

054

055

050

064

057

053

052

056

065

070

055

069

066

055

062

053

069

052

075

070

059

063

067

049

055

069

062

077

034

062

057

071

052

062

Parapurcellianatalia

sp

nov

050

046

050

051

054

054

046

046

079

060

056

066

072

063

064

055

062

060

057

058

046

068

053

063

063

069

042

063

045

054

073

050

062

073

056

054

060

048

054

Parap

urcelliaminuta

spn

ov

046

046

054

041

030

054

047

049

075

060

059

050

058

059

066

055

062

062

057

040

062

066

072

061

062

076

044

064

052

056

058

076

095

069

053

058

062

057

054

Parap

urcelliaperegrinator

058

058

058

050

049

054

046

054

071

063

049

056

070

062

060

055

062

060

057

070

053

063

073

077

079

056

052

078

049

061

058

057

064

079

052

048

066

055

055

Parapurcelliasilvicola

052

054

062

050

044

054

050

048

064

058

063

059

055

062

069

055

064

061

052

051

062

067

072

070

066

019

045

064

059

059

048

061

074

066

065

073

070

064

057

Parapurcelliamonticola

067

073

086

060

055

054

053

051

066

057

050

062

071

063

070

055

060

065

058

053

044

064

061

039

072

075

047

080

048

061

056

047

066

061

066

047

061

053

056

Parap

urcelliarumpian

a040

035

072

081

058

054

055

045

075

057

048

067

074

063

065

055

082

059

056

072

052

066

051

064

073

068

049

076

073

066

056

058

070

035

054

090

078

093

058

Parapurcelliafissa

060

057

065

048

059

054

057

060

063

057

049

056

066

063

060

055

062

061

051

064

056

065

062

038

074

051

057

079

050

059

043

048

069

061

067

051

071

055

060

Purcelliatransvaalica

064

068

060

051

053

054

056

057

062

055

044

054

065

055

069

055

077

068

052

054

044

068

069

066

064

058

060

058

060

071

050

043

054

051

073

055

054

056

058

Purcelliaillustrans

111

080

101

074

037

054

082

067

066

052

047

046

060

066

057

055

056

056

055

047

052

066

063

057

066

048

055

077

052

068

051

043

059

048

053

047

072

054

059

Purcelliagriswoldisp

nov

067

078

063

048

049

054

054

049

065

057

045

050

066

061

065

055

053

061

059

046

050

063

074

056

070

071

070

066

058

070

035

050

051

069

049

058

077

052

056

Purcellialawrenceisp

nov

071

081

067

052

046

054

056

052

089

054

046

040

054

061

066

055

059

059

069

047

038

074

067

054

072

068

056

056

056

065

051

050

050

050

042

062

068

052

057

Speleosiro

argasiform

is057

056

054

045

060

054

047

044

055

066

041

058

078

061

045

055

058

061

057

044

076

049

090

079

055

054

085

058

077

089

050

052

051

070

043

041

050

048

060

Rakaiaisolate

066

075

061

060

062

054

061

079

032

073

059

069

071

063

059

055

066

060

066

044

061

052

039

053

069

056

038

058

049

050

049

077

071

058

077

062

065

064

059

Rakaialin

dsayi

050

060

058

056

068

054

057

054

070

057

067

065

060

067

060

055

055

056

068

059

073

059

060

044

078

074

049

053

060

059

050

064

057

085

059

055

064

066

058

Rakaiamag

naau

stralis

060

047

055

055

055

054

046

052

059

078

064

074

072

066

054

055

061

064

073

060

060

064

050

056

070

047

058

068

062

052

053

074

049

049

058

061

070

077

058

Rakaiamedia

060

082

041

052

062

054

061

052

065

078

064

082

075

061

058

055

060

062

075

060

047

059

043

059

063

071

060

064

049

037

076

056

070

050

061

063

070

051

060

Rakaiasolitaria

065

071

056

052

051

054

060

066

068

067

074

075

063

069

053

055

078

056

065

060

053

064

047

055

063

051

057

081

050

048

074

072

056

065

075

069

073

052

058

Rakaiasorensenidigitata

076

071

066

060

034

054

087

052

064

060

049

030

039

084

055

055

065

065

065

060

068

057

072

072

072

051

067

069

061

062

061

078

076

078

077

073

071

070

048

Aorakicrypta

071

069

053

060

054

054

082

048

068

067

068

063

051

069

055

055

059

073

071

054

064

047

075

078

071

056

080

040

068

049

067

081

042

064

068

041

047

057

048

Aorakidenticula

075

068

054

061

068

054

085

055

052

057

068

055

048

077

054

055

070

060

056

065

047

073

071

049

074

066

053

055

057

035

085

061

070

067

058

040

059

054

060

Aorakiinerma

069

070

071

078

065

054

069

046

068

060

064

063

052

072

050

055

081

064

054

050

058

060

062

072

042

055

057

073

070

068

054

052

047

043

079

046

057

055

061

Aorakilong

itarsa

069

066

057

062

073

054

067

078

064

059

075

063

056

071

056

055

065

073

057

069

073

043

074

071

064

050

071

054

064

049

071

078

050

052

062

056

063

057

062

Aorakitumidata

066

062

067

047

063

054

073

062

066

055

074

087

071

070

065

055

051

065

072

062

095

023

074

082

079

081

077

063

082

060

064

105

068

085

055

059

071

069

069

Karripurcellia

harveyi

050

061

072

074

086

054

084

076

057

054

047

047

064

062

057

055

056

054

061

077

075

053

060

073

053

047

066

059

064

070

054

051

055

061

045

052

054

050

060

Karripurcellia

peckorum

046

050

066

077

087

054

067

056

050

067

045

047

062

064

056

055

059

059

074

071

069

056

061

059

063

045

055

049

082

067

054

055

042

054

081

071

051

090

055

Karripurcellia

sierwaldae

050

052

064

067

068

054

062

061

060

050

060

044

044

064

065

055

052

060

051

060

060

058

057

064

061

060

063

070

079

074

051

040

045

058

056

081

057

097

048

Austropurcelliascoparia

079

078

069

066

053

054

074

064

051

052

080

056

037

089

038

055

056

057

080

050

057

076

067

062

060

048

053

056

052

062

064

060

061

068

062

066

062

052

063

Austrop

urcelliawoo

dwardi

072

067

074

061

054

054

075

079

049

062

075

067

047

074

036

055

058

058

053

054

051

086

061

052

056

059

038

054

054

065

060

046

058

049

063

080

067

054

048

Chileogovea

jocasta

067

064

065

054

054

054

061

052

054

055

069

066

052

069

045

055

049

058

057

057

063

066

042

053

059

068

068

059

056

074

054

056

057

060

043

057

046

050

048

Chileog

ovea

oedipu

s064

068

070

070

048

054

070

076

081

056

070

056

046

069

056

055

054

059

054

085

072

060

047

044

076

057

066

055

072

074

048

064

054

076

046

052

041

060

048

Pettaluslampetid

es063

055

059

058

058

054

057

051

063

063

053

077

084

052

075

055

058

051

057

058

054

052

051

057

060

063

062

052

073

067

054

051

039

075

083

040

034

053

061

Pettaluscimicifo

rmis

063

061

050

062

048

062

060

058

044

065

044

054

079

048

070

055

042

048

069

049

060

051

057

051

058

062

082

054

051

061

077

059

060

053

072

046

062

051

057

Troglosiroaelleni

059

056

029

089

066

085

058

054

066

046

078

071

050

062

045

090

056

060

068

076

055

055

066

058

045

064

076

054

064

065

059

049

055

055

046

049

037

052

060

Troglosirolongifo

ssa

059

055

058

070

077

085

077

053

061

045

069

071

058

048

054

093

038

065

070

085

059

056

052

082

044

051

071

049

056

045

075

053

050

053

038

063

073

064

048

Troglosironinq

ua066

051

072

073

079

083

077

061

060

041

068

067

053

045

064

092

062

059

058

067

059

053

070

075

052

061

082

049

075

076

046

055

048

051

036

053

045

068

048

Huitaca

ventralis

045

045

040

050

059

085

046

058

056

041

049

070

083

032

075

081

031

044

049

051

062

061

070

030

046

062

071

090

108

096

085

062

043

073

043

063

033

084

048

Neogo

veasp

042

067

051

086

073

087

071

066

031

046

039

066

087

037

059

055

066

130

104

061

066

071

062

039

042

072

075

076

074

077

079

055

050

036

084

064

038

095

061

Metag

ovea

philipi

057

068

061

073

063

085

062

044

040

041

066

069

059

039

057

055

061

044

080

064

056

085

058

045

035

059

041

075

059

063

053

049

046

047

065

054

034

059

048

Cypho

phthalmus

duricorius

060

046

053

056

061

086

050

058

065

056

053

064

073

054

051

087

060

064

057

082

060

061

060

065

048

047

064

036

055

052

070

059

066

065

071

077

072

064

078

Parasirocoiffaiti

056

064

071

050

066

081

046

074

063

060

089

070

051

051

055

078

055

065

053

062

063

060

032

073

042

079

053

052

044

048

078

068

088

061

060

077

070

055

048

Siro

acaroides

052

040

038

053

081

085

049

077

058

060

057

079

073

055

059

096

032

061

049

068

045

071

040

057

055

059

058

043

065

064

063

065

066

050

059

067

056

059

048

Siro

rubens

063

050

064

053

063

085

050

069

056

049

076

069

054

060

051

087

073

067

078

089

056

064

034

060

046

084

064

038

056

054

068

064

076

049

045

063

068

058

073

Suzukielus

sauteri

064

047

063

048

073

054

047

101

069

053

063

050

048

052

075

055

060

062

067

077

059

058

042

069

068

069

052

050

049

048

075

061

066

056

069

065

074

050

068

Stylocellusramblae

060

072

062

053

058

054

062

073

041

098

074

042

040

035

071

055

025

044

038

053

105

053

072

051

030

061

082

052

056

052

067

075

059

072

063

066

048

047

099

Stylocellustambusisi

056

063

061

052

060

054

056

069

047

096

073

050

041

040

067

055

063

044

038

042

064

060

074

050

036

048

063

044

060

049

067

060

058

078

057

085

046

062

087

Fan

gensiscavernarus

047

044

040

056

075

054

046

070

036

085

050

042

058

041

084

055

077

044

038

066

076

035

052

064

049

055

070

046

051

044

056

067

062

071

050

058

060

057

096

Fan

gensisspelaeus

049

061

044

055

079

054

046

070

047

086

046

035

058

042

105

055

087

044

038

065

077

029

068

065

060

084

050

047

051

045

078

070

083

057

054

071

065

056

096

Systematic revision of South African pettalids Invertebrate Systematics 375

combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)

Methods and abbreviations

Specimens used in this study were provided by the followinginstitutions

AMNH American Museum of Natural History New York(USA)

BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard

University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand

Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo

Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg

(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg

Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)

Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg

(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen

Copenhagen (Denmark)

Specimen handling and illustration

In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was

photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy

Character coding

Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3

Phylogenetic analyses

The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)

The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the

376 Invertebrate Systematics B L de Bivort and G Giribet

continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data

Taxonomy

Order OPILIONES Sundevall

Suborder CYPHOPHTHALMI Simon

Family PETTALIDAE Simon

Genus Speleosiro Lawrence 1931

Diagnosis

Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII

Type species

Speleosiro argasiformis Lawrence 1931

Species included

Speleosiro argasiformis Lawrence 1931

Distribution

SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province

Remarks

This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement

Genus Purcellia Hansen amp Soslashrensen 1904

Diagnosis

Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)

Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)

Type species

Purcellia illustrans Hansen amp Soslashrensen 1904

Species included

Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov

Distribution

South Africa Limpopo Eastern Cape and Western CapeProvinces

Remarks

This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa

Purcellia griswoldi sp nov

(Figs 2 3 Table 3)

Material examined

Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985

Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981

Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981

Systematic revision of South African pettalids Invertebrate Systematics 377

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

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stra

ns

Cyp

ho

ph

tha

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s d

uri

cori

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Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

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oe

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us

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ita

ca v

en

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lis

Ch

ileo

gov

ea

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sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

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e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 4: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Tab

le2

Con

tinu

ouscharactermatrixas

itap

pearsin

the11

weigh

tedcombinedan

alysis

Character

definitio

nsaregivenin

App

endix3

12

34

56

78

910

1112

1314

1516

1718

1920

2122

2324

2526

2728

2930

3132

3334

3536

3738

39

Parap

urcelliastaregaisp

nov

054

059

058

052

053

053

059

047

064

051

058

063

047

067

069

075

048

071

070

056

055

049

062

073

057

057

052

052

054

066

054

068

048

048

056

056

067

061

052

Parap

urcelliaam

atolaesp

nov

047

061

057

056

056

064

052

047

065

057

053

072

051

062

061

059

062

070

066

056

042

063

062

053

053

058

058

057

072

055

050

061

043

049

056

056

066

062

049

Parapurcelliaconvexa

spn

ov

048

069

058

075

067

066

049

053

060

051

050

078

081

063

074

051

064

062

071

059

056

065

057

059

045

066

085

075

071

070

068

052

057

051

056

056

061

060

054

Parapurcellianatalia

sp

nov

044

056

057

052

053

057

049

047

057

049

060

063

048

050

071

061

105

082

074

060

053

059

061

076

059

059

052

052

065

056

043

065

043

047

056

056

071

058

049

Parapurcelliaminuta

spn

ov

042

056

056

052

053

046

050

047

057

054

081

075

047

068

072

082

048

069

062

060

059

069

059

050

052

057

052

052

043

067

055

064

057

052

056

056

067

058

048

Parapurcelliaperegrinator

059

061

058

059

055

057

059

047

077

055

065

065

048

063

049

048

066

061

057

061

055

056

057

062

071

059

060

055

058

061

053

062

053

047

056

056

065

059

059

Parapurcelliasilvicola

055

063

063

076

064

049

062

050

062

062

060

072

077

053

079

050

065

079

070

059

060

065

064

064

057

071

080

067

047

097

059

068

040

059

056

056

064

058

070

Parapurcelliamonticola

054

061

059

061

053

056

045

053

073

052

058

074

045

055

064

073

050

070

059

057

065

062

058

064

061

063

063

052

058

054

066

049

074

051

056

056

066

063

062

Parap

urcelliarumpiana

051

059

056

057

053

050

040

053

067

078

046

073

075

057

073

080

037

074

071

057

061

058

054

056

059

059

060

052

049

062

066

039

061

046

056

056

075

060

055

Parapurcelliafissa

043

064

064

052

053

056

059

053

065

052

055

070

071

069

076

063

055

066

060

057

063

079

078

060

051

085

052

052

062

036

068

083

053

053

056

056

068

060

052

Purcelliatransvaalica

069

064

057

052

053

048

060

058

061

058

041

071

070

057

074

050

061

053

072

050

067

050

056

054

056

056

052

052

046

056

054

051

050

036

056

056

074

058

048

Purcelliaillustrans

072

063

055

052

053

052

060

087

081

056

061

063

062

061

077

049

059

075

071

051

065

058

051

078

087

054

052

052

050

089

096

053

076

036

056

056

078

061

042

Purcelliagriswoldisp

nov

062

063

057

057

053

043

062

055

067

053

065

066

062

044

064

068

050

057

058

048

075

059

055

053

058

058

057

052

041

061

061

060

060

040

056

056

075

057

045

Purcellialawrenceisp

nov

062

061

056

057

053

051

066

057

057

053

059

069

067

047

054

066

053

068

067

048

080

063

052

056

050

055

057

052

050

063

068

064

068

039

056

056

075

053

044

Speleosiro

argasiform

is097

077

060

075

059

073

091

048

071

051

049

044

063

039

056

055

068

031

035

045

108

046

054

047

050

057

064

056

061

053

056

060

057

047

056

056

070

058

043

Rakaiaisolate

059

077

067

102

058

069

066

061

059

063

067

077

077

070

062

046

077

054

068

052

062

062

072

061

043

076

104

058

071

066

061

068

059

062

056

056

058

064

099

Rakaialin

dsayi

061

063

062

083

053

059

053

057

054

066

071

056

071

077

067

086

046

079

059

051

057

071

058

057

061

070

082

052

061

063

037

055

037

056

056

056

062

066

062

Rakaiamag

naau

stralis

074

068

064

078

061

085

070

047

077

085

072

052

071

057

043

053

067

068

062

054

062

062

066

068

058

060

072

059

078

065

065

059

067

053

056

056

065

058

075

Rakaiamedia

062

070

062

063

057

055

077

062

066

052

066

058

070

081

066

087

041

065

064

049

057

057

068

058

068

061

062

057

054

058

054

077

057

064

056

056

061

063

077

Rakaiasolitaria

057

065

059

071

053

065

052

060

079

053

074

080

051

055

051

046

070

065

068

053

077

043

055

037

060

065

073

052

069

069

067

055

065

066

056

056

053

062

074

Rakaiasorensenidigitata

057

071

055

054

055

075

071

080

065

066

056

093

052

085

038

054

064

069

065

051

075

075

053

066

069

054

055

056

083

060

059

080

058

074

056

056

047

053

078

Aorakicrypta

071

079

061

058

059

085

069

088

041

059

049

059

043

072

059

047

071

060

064

052

060

037

051

039

053

071

057

058

081

067

062

061

053

039

056

056

077

075

086

Aorakidenticula

057

067

055

068

062

063

048

080

054

054

056

065

051

085

058

060

067

068

071

053

066

065

051

057

069

055

071

064

066

059

063

050

063

044

075

072

071

070

070

Aorakiinerma

067

066

057

063

063

062

056

070

075

056

044

066

065

077

053

051

066

060

059

054

073

066

053

053

063

057

062

063

060

054

060

050

059

062

056

056

059

072

074

Aorakilong

itarsa

066

076

073

098

109

065

058

069

056

058

083

056

049

062

028

049

061

063

064

050

061

059

066

079

060

091

094

107

064

076

062

055

058

064

056

056

059

129

062

Aorakitumidata

064

072

058

081

069

079

070

072

063

068

071

073

079

083

054

080

060

073

070

050

062

072

055

054

062

060

083

070

081

085

062

070

050

070

075

096

080

059

071

Karripurcellia

harveyi

067

052

058

061

066

051

079

089

062

052

069

056

073

060

074

075

037

050

043

055

056

063

060

069

091

052

060

065

049

066

050

072

048

067

056

056

047

057

058

Karripurcellia

peckorum

068

062

057

071

068

069

062

070

051

094

057

064

070

057

052

045

071

046

037

055

060

064

057

059

076

054

069

067

064

070

046

053

037

068

056

056

047

056

056

Karripurcellia

sierwaldae

056

056

057

052

053

042

062

061

067

088

056

061

076

063

070

057

060

043

032

055

062

069

062

067

068

053

052

052

039

067

057

064

051

073

064

087

047

057

060

Austropurcelliascoparia

055

050

054

052

053

067

052

068

054

051

045

065

050

064

057

044

071

065

074

059

071

072

051

054

084

053

052

052

071

052

082

055

088

062

056

056

061

091

053

Austrop

urcelliawoo

dwardi

054

065

058

052

058

056

061

069

053

053

042

066

049

066

068

053

057

062

068

054

063

047

049

036

050

067

052

060

056

065

079

064

068

059

056

056

047

061

053

Chileogovea

jocasta

049

053

055

052

054

058

050

057

055

049

047

049

058

060

055

063

050

050

058

051

074

055

053

053

060

053

052

055

061

057

069

057

068

066

082

088

082

053

069

Chileog

ovea

Oedipus

070

053

054

052

053

064

070

074

057

062

053

052

043

064

054

051

058

072

078

053

071

012

049

043

054

052

052

052

061

052

063

061

070

062

092

091

083

051

048

Pettaluslampetid

es063

066

078

052

092

067

057

058

053

055

063

039

048

055

057

063

064

068

068

046

064

050

095

090

044

084

052

093

065

042

050

052

066

074

056

056

047

052

055

Pettaluscimicifo

rmis

100

110

140

066

095

089

082

076

066

058

074

033

039

021

057

058

058

056

046

041

056

085

125

064

044

116

059

074

070

039

047

055

071

068

056

056

047

050

061

Troglosiroaelleni

059

053

059

054

066

066

064

057

054

053

052

040

067

066

072

060

064

044

053

076

045

064

066

075

062

054

054

067

068

054

061

066

064

074

056

056

047

055

058

Troglosirolong

ifossa

051

045

055

052

073

064

048

070

049

060

057

042

057

053

069

043

069

053

064

079

041

059

055

061

066

052

052

081

070

049

049

052

055

074

056

056

047

056

053

Troglosironinq

ua059

045

057

053

086

068

049

074

049

066

048

044

074

054

064

052

063

048

059

076

044

054

063

072

068

052

054

092

069

062

076

048

070

074

056

056

047

057

072

Huitaca

ventralis

075

045

057

052

060

036

084

047

105

094

063

054

058

067

072

068

054

040

056

075

057

046

058

070

053

052

052

058

039

045

038

057

047

074

056

056

047

052

048

Neogo

veasp

067

056

058

065

070

031

070

075

081

099

042

055

067

047

063

068

042

031

045

074

050

038

061

073

061

052

063

069

033

035

035

055

044

074

056

056

047

050

045

Metagovea

philipi

045

045

054

052

053

041

045

057

065

055

056

056

066

051

053

054

056

048

062

078

057

057

053

050

031

052

052

052

036

042

056

048

073

074

056

056

047

053

068

Cypho

phthalmus

duricorius

050

045

055

052

053

058

041

050

039

060

067

044

044

042

027

050

072

058

065

078

039

065

057

071

065

052

052

052

062

059

067

044

072

074

056

056

047

056

065

Parasirocoiffaiti

044

045

055

054

055

058

056

047

050

052

079

050

041

060

056

080

051

064

061

086

042

064

060

062

058

052

056

057

068

053

052

081

063

068

056

056

055

053

064

Siro

acaroids

046

055

054

052

053

055

053

049

044

055

047

051

067

060

070

051

057

058

071

069

048

068

053

044

061

053

052

052

058

062

061

062

055

051

056

056

071

055

068

Siro

rubens

049

045

056

052

053

042

077

050

041

055

065

039

057

053

062

082

037

047

052

072

035

073

059

052

086

052

052

052

036

054

085

102

100

066

119

102

056

050

049

Suzukielus

sauteri

059

045

056

052

053

060

062

048

050

051

063

068

069

065

064

066

056

082

071

075

048

071

056

067

057

052

052

052

062

079

072

066

063

071

098

091

047

056

061

Stylocellusramblae

052

045

055

052

053

064

050

060

052

048

083

058

047

056

053

055

076

045

046

092

068

060

054

046

048

052

052

052

067

053

058

053

074

074

056

056

047

053

060

Stylocellustambusisi

074

045

057

052

053

066

049

059

046

067

058

052

050

041

029

059

076

047

056

082

066

060

057

070

056

052

052

052

061

047

055

041

065

074

056

056

047

052

062

Fan

gensiscavernarus

081

064

065

052

053

073

046

047

049

065

080

054

076

053

065

048

079

048

026

075

061

073

065

078

081

059

052

052

066

063

049

038

051

074

056

056

047

053

048

Fan

gensisspelaeus

073

045

057

052

053

078

093

047

050

061

093

065

077

056

064

076

053

068

030

076

054

069

056

069

066

052

052

052

076

068

055

086

061

074

056

056

047

057

054

374 Invertebrate Systematics B L de Bivort and G Giribet

4041

4243

4445

4647

4849

5051

5253

5455

5657

5859

6061

6263

6465

6667

6869

7071

7273

7475

7677

78

Parap

urcelliastaregaisp

nov

054

049

048

053

061

054

046

048

065

057

060

056

057

059

069

055

074

063

048

049

055

058

069

062

068

080

057

066

053

057

056

052

068

059

066

054

070

051

062

Parap

urcelliaam

atolaesp

nov

047

058

055

044

066

054

046

050

081

058

068

069

065

064

068

055

068

055

049

032

047

064

064

070

066

059

053

073

046

043

035

075

066

053

068

048

065

062

056

Parapurcelliaconvexa

spn

ov

036

030

068

095

053

054

055

050

064

057

053

052

056

065

070

055

069

066

055

062

053

069

052

075

070

059

063

067

049

055

069

062

077

034

062

057

071

052

062

Parapurcellianatalia

sp

nov

050

046

050

051

054

054

046

046

079

060

056

066

072

063

064

055

062

060

057

058

046

068

053

063

063

069

042

063

045

054

073

050

062

073

056

054

060

048

054

Parap

urcelliaminuta

spn

ov

046

046

054

041

030

054

047

049

075

060

059

050

058

059

066

055

062

062

057

040

062

066

072

061

062

076

044

064

052

056

058

076

095

069

053

058

062

057

054

Parap

urcelliaperegrinator

058

058

058

050

049

054

046

054

071

063

049

056

070

062

060

055

062

060

057

070

053

063

073

077

079

056

052

078

049

061

058

057

064

079

052

048

066

055

055

Parapurcelliasilvicola

052

054

062

050

044

054

050

048

064

058

063

059

055

062

069

055

064

061

052

051

062

067

072

070

066

019

045

064

059

059

048

061

074

066

065

073

070

064

057

Parapurcelliamonticola

067

073

086

060

055

054

053

051

066

057

050

062

071

063

070

055

060

065

058

053

044

064

061

039

072

075

047

080

048

061

056

047

066

061

066

047

061

053

056

Parap

urcelliarumpian

a040

035

072

081

058

054

055

045

075

057

048

067

074

063

065

055

082

059

056

072

052

066

051

064

073

068

049

076

073

066

056

058

070

035

054

090

078

093

058

Parapurcelliafissa

060

057

065

048

059

054

057

060

063

057

049

056

066

063

060

055

062

061

051

064

056

065

062

038

074

051

057

079

050

059

043

048

069

061

067

051

071

055

060

Purcelliatransvaalica

064

068

060

051

053

054

056

057

062

055

044

054

065

055

069

055

077

068

052

054

044

068

069

066

064

058

060

058

060

071

050

043

054

051

073

055

054

056

058

Purcelliaillustrans

111

080

101

074

037

054

082

067

066

052

047

046

060

066

057

055

056

056

055

047

052

066

063

057

066

048

055

077

052

068

051

043

059

048

053

047

072

054

059

Purcelliagriswoldisp

nov

067

078

063

048

049

054

054

049

065

057

045

050

066

061

065

055

053

061

059

046

050

063

074

056

070

071

070

066

058

070

035

050

051

069

049

058

077

052

056

Purcellialawrenceisp

nov

071

081

067

052

046

054

056

052

089

054

046

040

054

061

066

055

059

059

069

047

038

074

067

054

072

068

056

056

056

065

051

050

050

050

042

062

068

052

057

Speleosiro

argasiform

is057

056

054

045

060

054

047

044

055

066

041

058

078

061

045

055

058

061

057

044

076

049

090

079

055

054

085

058

077

089

050

052

051

070

043

041

050

048

060

Rakaiaisolate

066

075

061

060

062

054

061

079

032

073

059

069

071

063

059

055

066

060

066

044

061

052

039

053

069

056

038

058

049

050

049

077

071

058

077

062

065

064

059

Rakaialin

dsayi

050

060

058

056

068

054

057

054

070

057

067

065

060

067

060

055

055

056

068

059

073

059

060

044

078

074

049

053

060

059

050

064

057

085

059

055

064

066

058

Rakaiamag

naau

stralis

060

047

055

055

055

054

046

052

059

078

064

074

072

066

054

055

061

064

073

060

060

064

050

056

070

047

058

068

062

052

053

074

049

049

058

061

070

077

058

Rakaiamedia

060

082

041

052

062

054

061

052

065

078

064

082

075

061

058

055

060

062

075

060

047

059

043

059

063

071

060

064

049

037

076

056

070

050

061

063

070

051

060

Rakaiasolitaria

065

071

056

052

051

054

060

066

068

067

074

075

063

069

053

055

078

056

065

060

053

064

047

055

063

051

057

081

050

048

074

072

056

065

075

069

073

052

058

Rakaiasorensenidigitata

076

071

066

060

034

054

087

052

064

060

049

030

039

084

055

055

065

065

065

060

068

057

072

072

072

051

067

069

061

062

061

078

076

078

077

073

071

070

048

Aorakicrypta

071

069

053

060

054

054

082

048

068

067

068

063

051

069

055

055

059

073

071

054

064

047

075

078

071

056

080

040

068

049

067

081

042

064

068

041

047

057

048

Aorakidenticula

075

068

054

061

068

054

085

055

052

057

068

055

048

077

054

055

070

060

056

065

047

073

071

049

074

066

053

055

057

035

085

061

070

067

058

040

059

054

060

Aorakiinerma

069

070

071

078

065

054

069

046

068

060

064

063

052

072

050

055

081

064

054

050

058

060

062

072

042

055

057

073

070

068

054

052

047

043

079

046

057

055

061

Aorakilong

itarsa

069

066

057

062

073

054

067

078

064

059

075

063

056

071

056

055

065

073

057

069

073

043

074

071

064

050

071

054

064

049

071

078

050

052

062

056

063

057

062

Aorakitumidata

066

062

067

047

063

054

073

062

066

055

074

087

071

070

065

055

051

065

072

062

095

023

074

082

079

081

077

063

082

060

064

105

068

085

055

059

071

069

069

Karripurcellia

harveyi

050

061

072

074

086

054

084

076

057

054

047

047

064

062

057

055

056

054

061

077

075

053

060

073

053

047

066

059

064

070

054

051

055

061

045

052

054

050

060

Karripurcellia

peckorum

046

050

066

077

087

054

067

056

050

067

045

047

062

064

056

055

059

059

074

071

069

056

061

059

063

045

055

049

082

067

054

055

042

054

081

071

051

090

055

Karripurcellia

sierwaldae

050

052

064

067

068

054

062

061

060

050

060

044

044

064

065

055

052

060

051

060

060

058

057

064

061

060

063

070

079

074

051

040

045

058

056

081

057

097

048

Austropurcelliascoparia

079

078

069

066

053

054

074

064

051

052

080

056

037

089

038

055

056

057

080

050

057

076

067

062

060

048

053

056

052

062

064

060

061

068

062

066

062

052

063

Austrop

urcelliawoo

dwardi

072

067

074

061

054

054

075

079

049

062

075

067

047

074

036

055

058

058

053

054

051

086

061

052

056

059

038

054

054

065

060

046

058

049

063

080

067

054

048

Chileogovea

jocasta

067

064

065

054

054

054

061

052

054

055

069

066

052

069

045

055

049

058

057

057

063

066

042

053

059

068

068

059

056

074

054

056

057

060

043

057

046

050

048

Chileog

ovea

oedipu

s064

068

070

070

048

054

070

076

081

056

070

056

046

069

056

055

054

059

054

085

072

060

047

044

076

057

066

055

072

074

048

064

054

076

046

052

041

060

048

Pettaluslampetid

es063

055

059

058

058

054

057

051

063

063

053

077

084

052

075

055

058

051

057

058

054

052

051

057

060

063

062

052

073

067

054

051

039

075

083

040

034

053

061

Pettaluscimicifo

rmis

063

061

050

062

048

062

060

058

044

065

044

054

079

048

070

055

042

048

069

049

060

051

057

051

058

062

082

054

051

061

077

059

060

053

072

046

062

051

057

Troglosiroaelleni

059

056

029

089

066

085

058

054

066

046

078

071

050

062

045

090

056

060

068

076

055

055

066

058

045

064

076

054

064

065

059

049

055

055

046

049

037

052

060

Troglosirolongifo

ssa

059

055

058

070

077

085

077

053

061

045

069

071

058

048

054

093

038

065

070

085

059

056

052

082

044

051

071

049

056

045

075

053

050

053

038

063

073

064

048

Troglosironinq

ua066

051

072

073

079

083

077

061

060

041

068

067

053

045

064

092

062

059

058

067

059

053

070

075

052

061

082

049

075

076

046

055

048

051

036

053

045

068

048

Huitaca

ventralis

045

045

040

050

059

085

046

058

056

041

049

070

083

032

075

081

031

044

049

051

062

061

070

030

046

062

071

090

108

096

085

062

043

073

043

063

033

084

048

Neogo

veasp

042

067

051

086

073

087

071

066

031

046

039

066

087

037

059

055

066

130

104

061

066

071

062

039

042

072

075

076

074

077

079

055

050

036

084

064

038

095

061

Metag

ovea

philipi

057

068

061

073

063

085

062

044

040

041

066

069

059

039

057

055

061

044

080

064

056

085

058

045

035

059

041

075

059

063

053

049

046

047

065

054

034

059

048

Cypho

phthalmus

duricorius

060

046

053

056

061

086

050

058

065

056

053

064

073

054

051

087

060

064

057

082

060

061

060

065

048

047

064

036

055

052

070

059

066

065

071

077

072

064

078

Parasirocoiffaiti

056

064

071

050

066

081

046

074

063

060

089

070

051

051

055

078

055

065

053

062

063

060

032

073

042

079

053

052

044

048

078

068

088

061

060

077

070

055

048

Siro

acaroides

052

040

038

053

081

085

049

077

058

060

057

079

073

055

059

096

032

061

049

068

045

071

040

057

055

059

058

043

065

064

063

065

066

050

059

067

056

059

048

Siro

rubens

063

050

064

053

063

085

050

069

056

049

076

069

054

060

051

087

073

067

078

089

056

064

034

060

046

084

064

038

056

054

068

064

076

049

045

063

068

058

073

Suzukielus

sauteri

064

047

063

048

073

054

047

101

069

053

063

050

048

052

075

055

060

062

067

077

059

058

042

069

068

069

052

050

049

048

075

061

066

056

069

065

074

050

068

Stylocellusramblae

060

072

062

053

058

054

062

073

041

098

074

042

040

035

071

055

025

044

038

053

105

053

072

051

030

061

082

052

056

052

067

075

059

072

063

066

048

047

099

Stylocellustambusisi

056

063

061

052

060

054

056

069

047

096

073

050

041

040

067

055

063

044

038

042

064

060

074

050

036

048

063

044

060

049

067

060

058

078

057

085

046

062

087

Fan

gensiscavernarus

047

044

040

056

075

054

046

070

036

085

050

042

058

041

084

055

077

044

038

066

076

035

052

064

049

055

070

046

051

044

056

067

062

071

050

058

060

057

096

Fan

gensisspelaeus

049

061

044

055

079

054

046

070

047

086

046

035

058

042

105

055

087

044

038

065

077

029

068

065

060

084

050

047

051

045

078

070

083

057

054

071

065

056

096

Systematic revision of South African pettalids Invertebrate Systematics 375

combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)

Methods and abbreviations

Specimens used in this study were provided by the followinginstitutions

AMNH American Museum of Natural History New York(USA)

BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard

University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand

Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo

Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg

(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg

Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)

Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg

(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen

Copenhagen (Denmark)

Specimen handling and illustration

In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was

photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy

Character coding

Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3

Phylogenetic analyses

The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)

The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the

376 Invertebrate Systematics B L de Bivort and G Giribet

continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data

Taxonomy

Order OPILIONES Sundevall

Suborder CYPHOPHTHALMI Simon

Family PETTALIDAE Simon

Genus Speleosiro Lawrence 1931

Diagnosis

Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII

Type species

Speleosiro argasiformis Lawrence 1931

Species included

Speleosiro argasiformis Lawrence 1931

Distribution

SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province

Remarks

This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement

Genus Purcellia Hansen amp Soslashrensen 1904

Diagnosis

Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)

Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)

Type species

Purcellia illustrans Hansen amp Soslashrensen 1904

Species included

Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov

Distribution

South Africa Limpopo Eastern Cape and Western CapeProvinces

Remarks

This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa

Purcellia griswoldi sp nov

(Figs 2 3 Table 3)

Material examined

Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985

Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981

Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981

Systematic revision of South African pettalids Invertebrate Systematics 377

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 5: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

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060

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064

068

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Purcelliaillustrans

111

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067

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argasiform

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dsayi

050

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076

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075

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069

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069

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harveyi

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sierwaldae

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079

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jocasta

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oedipu

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veasp

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philipi

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phthalmus

duricorius

060

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Suzukielus

sauteri

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058

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050

068

Stylocellusramblae

060

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053

058

054

062

073

041

098

074

042

040

035

071

055

025

044

038

053

105

053

072

051

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048

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099

Stylocellustambusisi

056

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073

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038

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Fan

gensiscavernarus

047

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Fan

gensisspelaeus

049

061

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054

046

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046

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096

Systematic revision of South African pettalids Invertebrate Systematics 375

combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)

Methods and abbreviations

Specimens used in this study were provided by the followinginstitutions

AMNH American Museum of Natural History New York(USA)

BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard

University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand

Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo

Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg

(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg

Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)

Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg

(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen

Copenhagen (Denmark)

Specimen handling and illustration

In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was

photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy

Character coding

Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3

Phylogenetic analyses

The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)

The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the

376 Invertebrate Systematics B L de Bivort and G Giribet

continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data

Taxonomy

Order OPILIONES Sundevall

Suborder CYPHOPHTHALMI Simon

Family PETTALIDAE Simon

Genus Speleosiro Lawrence 1931

Diagnosis

Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII

Type species

Speleosiro argasiformis Lawrence 1931

Species included

Speleosiro argasiformis Lawrence 1931

Distribution

SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province

Remarks

This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement

Genus Purcellia Hansen amp Soslashrensen 1904

Diagnosis

Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)

Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)

Type species

Purcellia illustrans Hansen amp Soslashrensen 1904

Species included

Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov

Distribution

South Africa Limpopo Eastern Cape and Western CapeProvinces

Remarks

This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa

Purcellia griswoldi sp nov

(Figs 2 3 Table 3)

Material examined

Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985

Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981

Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981

Systematic revision of South African pettalids Invertebrate Systematics 377

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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Fig12

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398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 6: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

combining morphological and molecular data partitions (Boyerand Giribet 2007 Clouse et al 2009) Recently morphometricdata were shown to carry significant phylogenetic signalin Cyphophthalmi (Clouse et al 2009 de Bivort et al 2010)and Pettalidae specifically (de Bivort et al 2010) Thereforewe performed phylogenetic analyses of the South Africanpettalids as well as 15 outgroup species using both traditionaldiscrete morphological data and morphometric data (separatelyand in combination) Two phylogenetic hypotheses emergefrom these analyses one in which the South African pettalidsform a derived clade within Pettalidae (on their own or inconjunction with the Australian genus Austropurcellia) andthe other in which Parapurcellia is basal within Pettalidae andthe group of South African species is polyphyletic The latterhypothesis bears some similarities to earlier molecular studiesbut they had sparser taxon sampling with respect to the SouthAfrican pettalid fauna (Boyer and Giribet 2007 2009 Boyeret al 2007b)

Methods and abbreviations

Specimens used in this study were provided by the followinginstitutions

AMNH American Museum of Natural History New York(USA)

BMNH The Natural History Museum London (UK)CM Canterbury Museum Christchurch (New Zealand)FMNH Field Museum of Natural History Chicago (USA)MCZ Museum of Comparative Zoology Harvard

University Cambridge (USA)MHNG Museacuteum drsquohistore naturelle Genegraveve (Switzerland)MNHN Museacutee National drsquoHistoire Naturelle Paris (France)MNZ Te Papa TongarewaMuseum of New Zealand

Wellington (New Zealand)MZUSP Museu de Zoologia da Universidade de Satildeo Paulo

Satildeo Paulo (Brazil)NMSA Natal Museum of South Africa Pietermaritzburg

(South Africa)QM Queensland Museum Brisbane (Australia)SAM South African Museum Cape Town (South Africa)SMF Forschungsinstitut und Naturmuseum Senckenberg

Frankfurt am Main (Germany)USNM USA National Museum (Smithsonian Institution)

Washington DC (USA)WAM Western Australian Museum Perth (Australia)ZMH Zoological Museum of Hamburg Hamburg

(Germany)ZMB Museum fuumlr Naturkunde Berlin (Germany)ZMUC Zoologisk Museum University of Copenhagen

Copenhagen (Denmark)

Specimen handling and illustration

In total 35 pettalid and 15 outgroup cyphophthalmid taxawere analysed Animals from our collection and specimens onloan were preserved in ethanol (typically 95 in recentlycollected specimens 70 otherwise) and imaged usingscanning electron microscopy (SEM) and focus-stacked lightmicrographs according to the methods detailed in Clouse andGiribet (2007) For all descriptions the holotype male was

photographed under light microscopy when available andwith the lending institutionrsquos permission paratype males wereimaged by SEM and preserved on mounting stubs Tracings ofthe dorsal ventral and lateral body views and lateral appendageviews were generated by directly tracing over specimenmicrographs using Adobe Illustrator (Adobe Systems Inc SanJose CA) One male and one female paratype of Purcelliagriswoldi sp nov were dissected in ethanol and their genitaliamounted temporarily in glycerol for imaging by compoundmicroscopy

Character coding

Conspicuous discrete characters such as the position of theozophores were scored from our database of images Thestates of more subtle characters such as the ornamentation ofparticular appendage segments were confirmed by examinationunder light or scanning electron microscopy The discretecharacters used are described in Appendix 2 and enumeratedin Table 1 Continuous characters were scored from digitalimages of the dorsal ventral and lateral views of the animalsusing the software ImageJ (Research Services Branch NIMHNIH Bethesda MD) as detailed in de Bivort et al (2010)Raw measurements and scaled shape descriptor characterswere analysed for pair-wise independence and when found tobe dependent collapsed using principal components analysisbased on an independence cutoff of 131 which was previouslyfound (deBivort et al 2010) to yield the treesmost consistentwithprevious studies These methods were performed using customscripts in MATLAB (Mathworks Inc Natick MA) and yieldedthe 78 simple and complex continuous characters detailed inTable 2 and Appendix 3

Phylogenetic analyses

The discrete character matrix was analysed in the phylogeneticprogram Tree analysis with New Technology (TNT) (Goloboffet al 2008) For the parsimony analysis under equal weightingheuristic searches were performed using the lsquoNew Technologyrsquooption and 1000 random addition sequences In all tree searchingthe Sectorial Search (Goloboff 1999) Ratchet (Nixon 1999)Drift (Goloboff 1999) and Tree Fusing (Goloboff 1999) optionswere enabled Under implied weighting (Goloboff 1993) for kranging from 1 to 6 300 random addition replicates were usedBootstrapping was done over 100 replicates each using 300random addition sequences and the absolute node frequencieswere recorded The parameters used for the continuous datasetwere identical with the exception of the bootstrap analysis inwhichonly100 randomaddition sequenceswereusedbecause thecontinuous search runs were considerably slower All resultingtrees were rooted with Stylocellidae as an outgroup followingBoyer et al (2007b)

The relative weighting of data partitions in combined analysescan influence tree topology Precise relative weighting could beachieved by iteratively weighting a partition solving for treeswith that dataset determining the length of the tree contributedby the weighted characters adjusting their weight by their newlength contribution etc until the tree and data partition weightsconverged Alternatively approximate relative weighting(Clouse et al 2009) can be accomplished by multiplying the

376 Invertebrate Systematics B L de Bivort and G Giribet

continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data

Taxonomy

Order OPILIONES Sundevall

Suborder CYPHOPHTHALMI Simon

Family PETTALIDAE Simon

Genus Speleosiro Lawrence 1931

Diagnosis

Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII

Type species

Speleosiro argasiformis Lawrence 1931

Species included

Speleosiro argasiformis Lawrence 1931

Distribution

SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province

Remarks

This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement

Genus Purcellia Hansen amp Soslashrensen 1904

Diagnosis

Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)

Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)

Type species

Purcellia illustrans Hansen amp Soslashrensen 1904

Species included

Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov

Distribution

South Africa Limpopo Eastern Cape and Western CapeProvinces

Remarks

This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa

Purcellia griswoldi sp nov

(Figs 2 3 Table 3)

Material examined

Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985

Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981

Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981

Systematic revision of South African pettalids Invertebrate Systematics 377

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

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underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

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4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

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Page 7: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

continuous partition of the combined dataset (in its original form)by the factoraLDLCwherea is thefinal desired contributionofthe continuous data portion compared to the discrete portion (ie18th 14th etc) LC is the length of the tree inferred from thecontinuous data in their original form and LD is the length of thetree inferred from the discrete data

Taxonomy

Order OPILIONES Sundevall

Suborder CYPHOPHTHALMI Simon

Family PETTALIDAE Simon

Genus Speleosiro Lawrence 1931

Diagnosis

Ozophores in a completely dorsal position Eyes and eye lensesabsent Male coxae IV endites lacking anterior projectionsDentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position Male anal platebilobed with a large central depression spanning roughly halfthe area of the anal plate Scopulae originating on the anteriorventral surface of the anal plate along the edges of the raisedlateral portions Anal gland on tergite VIII

Type species

Speleosiro argasiformis Lawrence 1931

Species included

Speleosiro argasiformis Lawrence 1931

Distribution

SouthAfrica only known from the type localityWynberg Cavesin Table Mountain Cape Peninsula Western Cape Province

Remarks

This genus is monotypic and nests within the genus Purcelliain some of our phylogenetic analyses (see below) consistentwith its type locality falling within the range of that genusMorphologically however it exhibits strong troglomorphicmodifications to both limbs and body that justify its retentionas a separate genus in the absence of consistent phylogeneticplacement within Purcellia Analysis of molecular data for thisspecies in the future may help resolve its specific phylogeneticplacement

Genus Purcellia Hansen amp Soslashrensen 1904

Diagnosis

Ozophores in a completely dorsal position Eyes presentincorporated into the base of the ozophores without lensesMale coxae IV endites lacking anterior projections (as inFig 2I) Dentition of the mobile digit of the chelicerae non-uniformwith the largest tooth in a central position (as in Fig 2N)

Male anal plate bilobed with a large central depression spanningroughly half the area of the anal plate (as in Fig 2K L) Scopulaeoriginating on the anterior ventral surface of the anal plate alongthe edges of the raised lateral portions Anal gland on tergite IXwith its opening oriented ventrally (as in Fig 2L)

Type species

Purcellia illustrans Hansen amp Soslashrensen 1904

Species included

Purcellia illustrans Hansen amp Soslashrensen 1904 Purcelliatransvaalica Lawrence 1963 Purcellia griswoldi sp novPurcellia lawrencei sp nov

Distribution

South Africa Limpopo Eastern Cape and Western CapeProvinces

Remarks

This genus includes most members of the genus Purcellia afterRosas Costa (1950 Giribet 2000) except Parapurcelliaperegrinator new comb which is here transferred toParapurcellia The species of Purcellia have large depressionson the male anal plate and no projections in the gonostome walland are distributed in southern SouthAfricawith the exception ofPurcellia transvaalica found in north-eastern South Africa

Purcellia griswoldi sp nov

(Figs 2 3 Table 3)

Material examined

Holotype lt (NMSA) from indigenous forest in Diepwalle For Sta(33570S 23100E 548m [1 800 ft]) 22 km NE of Knysna Cape ProvinceSouth Africa leg C Griswold J Doyen T M Griswold 11ndash13xi1985

Paratypes 3 lt 5 (NMSA) same collecting data as holotype 1 lt(MCZ)mounted for SEM 1lt 1 (AMNH158) same locality as holotypeleg S Peck J Peck 12xii1981 8lt 9 (FMNH81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 lt (AMNH 177) fromBerlese funnel of forest and log litter from Stormsrivier State Forest nearGoesabos Eastern Cape Province South Africa leg S Peck J Peck20xii1981 2 lt 2 (AMNH 192) from Berlese funnel of forest and loglitter from Goudveld forest near Knysna Cape Province South Africa legS Peck J Peck 28xii1981 21 lt 26 (FMNH 81ndash621) from forest litterfrom Tzitzikama Forest National Park Eastern Cape Province South Africaleg S Peck 20xii1981

Additional material examined South Africa Cape Province 3juveniles (NMSA) same collecting data as holotype 2 lt 1 juvenile(FMNH 81ndash603) from elephant dung same locality as holotype legS Peck 12xii1981 13 juveniles (FMNH 81ndash604) from forest littersame locality as holotype leg S Peck 13xii1981 1 juvenile (AMNH192) from Berlese funnel of forest and log litter from Goudveld forest nearKnysna legSPeck JPeck28xii1981 19lt 9 3 juveniles (AMNH) fromBerlese funnel of forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck J Peck 20xii1981 21 lt 16 59 juveniles(FMNH 81ndash621) from forest litter from Tzitzikama Forest National ParkEastern Cape Province leg S Peck 20xii1981 48 juveniles (FMNH81ndash621) from forest litter from Tzitzikama Forest National Park EasternCape Province leg S Peck 20xii1981 1 juvenile (FMNH 81ndash732) fromforest litter from Goudveld forest near Knysna leg S Peck 28xii1981

Systematic revision of South African pettalids Invertebrate Systematics 377

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 8: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

A B

E

F G H

I

F G H

I

N O P

Q

N O P

R S TR S TQ

J K L

M

J K L

M

DC

378 Invertebrate Systematics B L de Bivort and G Giribet

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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Fig12

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ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 9: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Diagnosis

With a relatively short unenlarged tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Lateral margins of coxae IV visible along almost theirentire length in the dorsal view

Description

Male

Total length 243mm width across ozophores 083mmgreatest width 149mm in the opisthosoma nearly as wide inthe prosoma behind the ozophores greatest height 105mmlengthndashwidth ratio 175 Body reddish-brown (when preserved in70 ethanol) Anterior margin of dorsal scutum concave withoutlateral projections prosoma roundly triangular Eyes presentvisible as a white mass under ozophores lenses absent Dorsalvertical-facing ozophores with circular opening ornamentationuniform and non-directional (Fig 2A C) Transverse prosomalsulcus present but inconspicuous Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous Coxae IV wide with lateral marginsentirely visible beyond the lateral prosomal margin in thedorsal view Dorsal scutum flat maximum width and height atopisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites Coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 2A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae III notrunning along the margin between coxae II and III coxae IV

endites running parallel to coxae IV midline suture for a distanceslightly longer than the gonostome Coxal endites formingsmooth sternal plate with greatest width 034mm betweencoxae III and IV Pores of coxal glands visible at innermargins between coxae III and IV Sternum absent Coxae IVendites without projections Gonostome roundly semi-circular0091mm long and 015mm wide delimited by coxae IVeverywhere except posterior margin Lateral walls formed byvery slightly elevated endites of coxae IV (Fig 2I) Coxae IVwithgranulated ridges flanking gonopore and coxae IV endites

Spiracles circular slightly open to the lateral posterior withmaximum diameter 011mm (Fig 2J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7curved anteriorly through the midline Sternites 8 and 9 andtergite IXfree not formingacoronaanalis (Fig 2DK)Tergite IXuniform in length bilobed with a single central opening of theanal gland oriented ventrally tergite VIII lacking opening of theanal glands Anal plate measuring 021 030mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 2D L) Cuticle withtubercularndashmicrogranulate morphology (Murphree 1988) in allventral areas including coxae and anal plate except on coxal andpalpal enditesmargin of ventral surface between the sternites andtergites and the depression of the anal plate (Fig 2D IndashL)

Chelicerae relatively elongate with few setae first articlewith microgranulation on dorsal and lateral surfaces secondarticle appearing without conspicuous ornamentation underlight microscopy Proximal article without prominent dorsaland ventral processes (Fig 2M) Second article sub-cylindricalits widest portion near middle of article but closer to articulationwith mobile digit bearing a longitudinal apodeme on the lateral

A B

Fig 3 Purcellia griswoldi sp nov (A) Spermatopositor of male paratype Scale bar 01mm (B) Ovipositorof female paratype Scale bar 02mm

Fig 2 Purcellia griswoldi sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashT) Scanning electronmicrographs (SEMs)ofmale paratype (I) ventral gonostome complex (J) spiracle (K) anal region arrowhead indicatesopening of the anal gland (L) magnified view of anal region (M) chelicer (N) chelicer dentition (O) palp arrowhead indicates ventral process on trochanter(P) palpal claw (Q) tibia andmetatarsus of leg II (R) leg II claw (S) leg IV tarsus (T) magnified view of leg IV tarsus showing adenostyle Scale barsAB 1mmCndashH 1mm I 02mm J 005mm K L 02mm M 05mm N 01mm O 05mm P 005mm Q 02mm R 01mm S 02mm T 01mm

Systematic revision of South African pettalids Invertebrate Systematics 379

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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us

Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 10: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

side Proximal article 059mm long 024mm wide secondarticle 085mm long 015mm wide moveable finger 027mmlong 0052mm wide Dentition on mobile digit non-uniformwith eight small denticles on the proximal portion of the digit andfive larger distal denticles Fixed digit with 13 uniform denticles(Fig 2N)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 2O) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0041mm long

Legs (Fig 2B Table 3) with all claws smooth long and hook-like without lateral ornamentation (Fig 2R S) Surfaces of all

trochanters femurs patellae tibae and metatarsi of legs III andIV entirely ornamented Metatarsi of legs I and II ornamentedproximally and smooth distally All tarsi appearing smooth bylight microscopy ornamented sparsely by brown granulesTarsus of leg I lacking distinct solea Tarsus IV bisegmented(Fig 2ST) carrying a lamelliformadenostyle towards themiddleof the proximal segment approximate length of adenostyle014mm bearing no setae (Fig 2T) distal margin at 44 oftarsal length

Spermatopositor short (035mm) typical of pettalids(Fig 3A) Setal formula 4 6 44 Dorsal side of penis with agroupof four longmicrotrichiae on each sidewith bases arranged

Table 3 Lengths of the segments of the legs and palps of the new species described hereinLengthswidths of legs (L) and palps (P) in mm Parentheses indicate length-to-width ratio Tarsus II indicates the portion of the tarsus distal to the dividing suture

Trochanter Femur Patella Tibia Metatarsus Tarsus Tarsus II Sum

Parapurcellia amatolaeP 237101 (235) 35195 (369) 26682 (324) 29379 (371) 33779 (427) 1484LI 186169 (110) 596193 (308) 313172 (182) 405168 (241) 269141 (190) 480153 (314) 2248LII 191155 (123) 497162 (307) 280174 (160) 341173 (197) 245135 (181) 427149 (286) 1979LIII 189156 (121) 356159 (224) 267170 (157) 297166 (179) 241120 (200) 323117 (276) 1673LIV 262158 (166) 535166 (322) 307183 (168) 333171 (195) 259132 (196) 395164 (241) 197124 (159) 2092

Parapurcellia staregaiP 19194 (203) 30581 (377) 22983 (276) 24765 (380) 28376 (372) 1255LI 162131 (123) 487139 (350) 256142 (180) 327139 (236) 230128 (181) 396147 (270) 1858LII 163131 (124) 404135 (300) 230140 (165) 272136 (201) 201129 (156) 355133 (267) 1626LIII 166125 (133) 327125 (262) 210142 (148) 223134 (167) 18599 (187) 285102 (280) 1397LIV 236126 (187) 441143 (307) 235144 (164) 297139 (213) 197118 (166) 350131 (267) 15196 (158) 1756

Parapurcellia convexaP 24175 (321) 34391 (377) 22685 (266) 25575 (340) 27276 (358) 1337LI 159149 (107) 500158 (316) 261155 (168) 324155 (209) 185127 (145) 381159 (240) 1810LII 153127 (121) 415141 (294) 203150 (135) 267151 (177) 174120 (145) 302136 (222) 1514LIII 150138 (108) 323141 (229) 199142 (140) 237142 (167) 201105 (192) 282105 (267) 1392LIV 245135 (182) 394149 (264) 244153 (160) 288168 (171) 189134 (141) 336175 (192) 165110 (150) 1696

Parapurcellia nataliaP 15398 (156) 36786 (427) 23975 (319) 29878 (382) 30074 (405) 1357LI 160156 (103) 504149 (339) 244147 (166) 338142 (239) 238132 (181) 434146 (298) 1918LII 170138 (123) 427130 (328) 224134 (167) 279141 (197) 208116 (180) 375123 (304) 1683LIII 169138 (122) 347134 (259) 199142 (140) 252134 (188) 198102 (195) 30791 (339) 1472LIV 232137 (170) 423144 (294) 216152 (143) 289157 (184) 210117 (180) 352128 (275) 16697 (172) 1722

Parapurcellia minutaP 18770 (267) 28267 (421) 19863 (314) 21453 (404) 21956 (391) 1100LI 130102 (127) 417119 (351) 195124 (157) 244117 (208) 17099 (171) 317111 (284) 1473LII 145101 (143) 323111 (292) 156127 (123) 208118 (176) 14598 (148) 262110 (237) 1238LIII 128104 (124) 26299 (264) 124104 (119) 175106 (165) 13980 (174) 22176 (291) 1050LIV 160106 (151) 353114 (308) 196135 (145) 217118 (183) 158101 (156) 261115 (226) 11378 (144) 1344

Purcellia griswoldiiP 272120 (227) 51098 (520) 37895 (398) 41380 (516) 38592 (418) 1958LI 215237 (091) 779202 (386) 371200 (186) 544185 (295) 304165 (185) 633175 (361) 2848LII 241268 (090) 766230 (333) 376242 (156) 527240 (219) 329171 (192) 639162 (396) 2878LIII 198220 (090) 503180 (280) 273190 (143) 380190 (200) 236141 (167) 455120 (378) 2046LIV 286220 (130) 736217 (340) 331214 (155) 487216 (226) 264192 (137) 534209 (256) 294139 (212) 2637

Purcellia lawrenceiP 281120 (234) 486109 (446) 37395 (393) 39388 (447) 40592 (440) 1938LI 214203 (106) 738200 (369) 345192 (179) 552188 (293) 303153 (198) 626179 (351) 2777LII 204201 (102) 590188 (314) 270190 (142) 448191 (235) 242148 (164) 519157 (331) 2273LIII 165205 (081) 536184 (291) 283194 (146) 416191 (218) 252137 (184) 478143 (335) 2132LIV 281201 (140) 718231 (311) 374206 (181) 493224 (220) 330193 (171) 583227 (257) 306145 (211) 2779

380 Invertebrate Systematics B L de Bivort and G Giribet

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

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F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 11: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

in a V and not fused Distal margin of medium lobe of thespermatopositor bilobed with three short apical microtrichiaeon each lobe Ventral side with two long microtrichiae on eachside the bases arranged as a parallelogram not fused at the baseGonopore complexwith two shortmovablefingers in the shape ofcurvedhooks slightly surpassing the edgeof the spermatopositor

Female

Similar to male in non-sexual characters (Fig 2FndashH) Femaleparatype slightly larger than male holotype Total length255mm width across ozophores 083mm greatest width140mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 105mm length-width ratio 183 Anal plate in a terminal position compared withmale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Ovipositor not sheathed at base longer than 1mm when notextendedwith bilobed terminal segment (Fig 3B) Each segmentwith a row of six setae Those in the second and third to lastsegments more than twice as long as the remainder Apical lobeselongated with numerous short setae each lobe with a singlelong apical seta and a subapical lateral multibranched sensoryprocess

Remarks

This species is likely sympatric with Purcellia lawrencei and hasbeen found abundantly at several locations being along withP illustrans one of the most broadly distributed South Africanpettalids This is unusual within South African Cyphophthalmimost species of which are found at few if any localities beyondthe type locality

Distribution

Known from the Eastern and Western Cape Provinces of SouthAfrica from Goudveld Forest in the west to Tzitzikama Forest inthe east

Habitat

Specimens have been collected using Berlese funnelling offorest and log litter and directly off elephant dung in theKnysna subtropical moist broadleaf forest

Etymology

This species is named after Charles E Griswold an arachnologistcolleague who collected the type specimens and who hascontributed enormously to the knowledge of Afromontanearachnids

Purcellia lawrencei sp nov

(Fig 4 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo1875-KnysnaWGR Jan 1939rsquo Records at the NMSA indicate the

following collected nearKnysna (34020S 23020E)WesternCape ProvinceSouth Africa leg W G Rump i1939

Paratype 1 (NMSA) same collecting data as holotype

Diagnosis

Lacking a longitudinally expanded tergite IX Anterior margin ofdorsal prosoma at the site of cheliceral articulation not projectinglaterally Prosoma wider than opisthosoma None of the lateralmargin of coxae IV visible beyond the lateral margins of theprosoma in dorsal view

Description

Male

Total length 243mm width across ozophores 085mmgreatest width 139mm in the prosoma behind the ozophoresgreatest height 104mm length-width ratio 175 Body orange-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes present visible as a white mass underozophores lenses absent Dorsal vertical-facing ozophores withcircular opening ornamentation uniform and non-directional(Fig 4A C K) Transverse prosomal sulcus present butinconspicuous (Fig 4A C) Transverse opisthosomal sulcivisible Mid-dorsal longitudinal opisthosomal sulcus presentbut inconspicuous (Fig 4C) Dorsal scutum flat maximumwidth in prosoma maximum height at opisthosomal area

Coxae of legs I II and III free Ventral prosomal complexwith coxae I not meeting at the midline separated by the palpalendites Coxae II III and IV meeting at the midline marginbetween coxae II and III and margin between coxae III and IVboth reaching themidline (Fig 4AD I) near each other Anteriormargin of coxae II endites enlarged curving laterally andposteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures giving coxae III endites av-shaped appearance coxae IV endites running parallel tocoxae IV midline suture for a distance slightly longer than thegonostome Coxal endites forming a smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae IIIand IV Sternum absent Coxae IV endites without projectionsGonostome round to semi-circular 0083mm long and 013mmwide delimited by coxae IV everywhere except posteriormargin Lateral walls formed by very slightly elevated enditesof coxae IV (Fig 4I) Coxae IV with ornamented ridges flankinggonopore and coxae IV endites

Spiracles circular slightly open to the posterior withmaximum diameter 0096mm (Fig 4J) Sternal opisthosomalglands absent Sternite 8 and posterior margin of sternite 7 curvedanteriorly through the midline Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 4A L) Tergite IX uniformin length not bilobed with a single central opening of the analgland oriented ventrally tergite VIII lacking opening of the analglands Anal plate measuring 023 031mm with wideanterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along the margins of the raisedlateral portions of the anal plate (Fig 4) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites

Systematic revision of South African pettalids Invertebrate Systematics 381

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 12: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

AP

sc

opTIX

S9 S8S7

S6

TVIII

S5

C

I J K L

A B

D E

CIV

S1

G

CI

CII

CIII

EP

ECIII

ECIVCG

ECI

ECII

F G HF G H

382 Invertebrate Systematics B L de Bivort and G Giribet

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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Fig12

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k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 13: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

margin of ventral surface between the sternites and tergites andthe depression of the anal plate (Fig 4C D IndashL)

Chelicerae relatively robust with few setae first articlewith microornamentation on dorsal and lateral surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with small dorsalprocess lacking ventral processes (Fig 4B) Second articlesub-cylindrical its widest portion through middle of articlebearing a longitudinal apodeme on the lateral side Proximalarticle 069mm long 030mm wide second article 095mmlong 018mm wide moveable finger 028mm long 0075mmwide Dentition on mobile digit non-uniform with 8 smalldenticles on the proximal portion of the digit and 4 largerdistal denticles

Palp with club-shaped trochanter narrowing posteriorly(Fig 4B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0035mm long

Legs (Fig 4B Table 3) with all claws smooth long and hook-like without lateral ornamentation Surfaces of all trochantersfemurs patellae tibae and metatarsi of legs III and IV entirelyornamentedMetatarsi of legs I and II ornamented proximally andsmooth distally Tarsi of legs I and II ornamented by sparse browngranules Tarsi of legs III and IV appearing smooth by lightmicroscopy Tarsus of leg I lacking distinct solea Tarsus IVbisegmented (Fig 4B) carrying a lamelliform adenostyletowards the middle of the proximal segment approximatelength of adenostyle 0096mm bearing no setae distal marginat 43 of tarsal length

Spermatopositor not studied due to the single male specimenavailable (the holotype)

Female

Similar to male in non-sexual characters (Fig 4FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 260mm width across ozophores 081mm greatest width141mm in the opisthosoma nearly as wide (135mm) in theprosoma behind the ozophores greatest height 108mm length-width ratio 193 Anal plate in a terminal position compared tomale though not visible from the dorsal view without a bilobedelevated anteriormargin TergiteVIII not bilobed Tarsus IV thinlonger than inmales not bisegmentedGonostome area typical ofpettalids with coxae of leg IV not meeting in the midline

Remarks

This species is likely sympatric with Purcellia griswoldi

Distribution

Only known from the type locality near Knysna Western CapeProvince South Africa

Habitat

No further information is available beyond the collection localitywhich is within the Knysna subtropical moist broadleaf forest

Etymology

The species is named after Reginald Frederick Lawrence aprolific South African zoologist who published more than 200articles mostly focusing on South African arthropods

Genus Parapurcellia Rosas Costa 1950

Diagnosis

Ozophores slightly elevated above carapace facing 45 (as inFig 5E) Eyes and eye lenses absent Male coxae IV enditesbearing anterior projections in the gonostomewall (as in Fig 5F)Dentition of the mobile digit of the chelicerae uniform (except inthe case of Parapurcellia peregrinator (Lawrence 1963) newcombination) Male anal plate bilobed with a small centraldepression spanning less than one third the area of the analplate (as in Fig 5I) Scopulae absent or originating on thecentral ventral surface or posterior margin of the anal platetypically along the edges of the raised lateral portions Analgland on tergite IX with its opening oriented posteriorly (as inFig 5I)

Type species

Parapurcellia fissa (Lawrence 1939)

Species included

Parapurcellia fissa (Lawrence 1939) Parapurcellia monticola(Lawrence 1939) Parapurcellia rumpiana (Lawrence 1933)Parapurcellia silvicola (Lawrence 1939) Parapurcelliaperegrinator (Lawrence 1963) new combinationParapurcellia amatola sp nov Parapurcellia convexa spnov Parapurcellia minuta sp nov Parapurcellia nataliasp nov Parapurcellia staregai sp nov

Distribution

South Africa Eastern Cape Kwazulu-Natal and MpumalangaProvinces

Remarks

This genus includes all members of the old genus Parapurcellia(sensu Rosas Costa 1950 Giribet 2000) plus Purcelliaperegrinator Lawrence 1963 all animals with smalldepressions on the male anal plate and anterior projections inthe gonostomewall They are distributed from southern to easternSouth Africa

Fig 4 Purcellia lawrencei sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashL) Automontage photographs of various holotype body parts For all species in which there were insufficient specimens to SEM paratypesautomontage photographs of the holotype are provided along with annotated tracings (below) to clarify their details Specific morphological features are asfollows (I) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP is the palpal endite CG coxalgland G gonostome and S1 sternite 1 (J) Spiracle (K) Ozophore Dashed line indicates eye incorporated into base of ozophore (L) Anal region S5ndash9 indicatesternites 5ndash9AP anal plate SC scopulae on anal plate TIX tergite IX TVIII tergiteVIII andOPopening of the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale bars A B 1mm CndashH 1mm IndashL 01mm

Systematic revision of South African pettalids Invertebrate Systematics 383

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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99 51 69

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Fig12

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onyusingcombineddiscreteandcontinuous

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hesearch

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eights(EW)and

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urcelliaisfoun

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11

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ithinthePettalid

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ata

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umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 14: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

C

A B

D E

F G H IF G H I

N O P QN O P Q

J K L MJ K L M

384 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 15: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Parapurcellia amatola sp nov

(Fig 5 Table 3)

Material examined

Holotype lt (NMSA) from Amatola Mountains near HogsbackEastern Cape Province South Africa leg D Brolhers iv1965

Paratypes 2 lt (NMSA) same collecting data as holotypeAdditional material examined 1 juvenile (NMSA) same collecting

data as holotype

Diagnosis

Tergite VII entirely convex Tergite VIII not prominently lobedAnterior margin of gonopore rounded Mid-dorsal longitudinalopisthosomal sulcus absent Margins between coxae III and IVmeeting in an acute angle at the midline

Description

Male

Total length 209mm width across ozophores 086mmgreatest width 120 in the prosoma behind the ozophoresgreatest height 079mm length-width ratio 174 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly trapezoidal Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 5A C H)Transverse prosomal sulcus present but inconspicuous mostprominent laterally (Fig 5A C) Transverse opisthosomal sulciinconspicuous Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum flat maximum width and height inprosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV reaching the midline (Fig 5A D F) Endites of coxae IIand III running along their margins giving coxae III endites a v-shaped appearance coxae IV endites running parallel to coxae IVmidline suture for a distance longer than the gonostome Coxalendites forming smooth sternal platewith greatest width 043mmbetween coxae III and IV Pores of coxal glands visible at innermargin between coxae III and IV Sternum absent Coxae IVendites with horn-like projections on anterior margin ofgonostome Gonostome elliptical 0058mm long and 012mmwide and delimited anteriorly by coxae IV Lateral walls formedby elevated endites of coxae IV (Fig 5F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 5G) Sternal opisthosomal glandsabsent Sternites 7 and 8 narrow curved anteriorly through themidline posterior margin of sternite 6 similarly curved Sternites8 and 9 and tergite IX free not forming a corona analis (Fig 5I)

Tergite IX and posterior margin of tergite VIII curving to theanterior at the mid-line appearing w-shaped Tergite VIIIlacking anal glands tergite IX with a single central openingof the anal gland oriented posteriorly Anal plate measuring016 025mm with an ungranulated anterior-pointingv-shaped depression and scopulae originating in the centralventral surface (Fig 5I) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites anddepression of anal plate (Fig 5FndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with prominent dorsal crestlacking ventral processes (Fig 5B J) Second article robustsub-cylindrical its widest portion near articulation with mobiledigit bearing a longitudinal apodemeon the lateral side Proximalarticle 054mm long 025mm wide second article 073mmlong 019mm wide moveable finger 019mm long 0058mmwide Dentition on mobile digit non-uniform with 6 smalldenticles on the proximal portion and 4 larger distal denticles11 uniform denticles on fixed digit

Palp with club-shaped trochanter narrowing posteriorly(Fig 5B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 039mm long

Legs (Fig 5B NndashQ Table 3) with all claws smooth long andhook-like lacking lateral pegs or other ornamentation (Fig 5OP) Surfaces of all trochanters femurs patellae and tibae entirelyornamented metatarsi partially ornamented (Fig 5N) Tarsus ofleg I lacking a distinct solea All tarsi appearing smooth by lightmicroscopy Tarsus IV bisegmented (Fig 5B P Q) carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length of adenostyle 010mm bearingno setae (Fig 5Q) distal margin at 49 of tarsal length

Spermatopositor not studied

Female

Unknown

Distribution

Only known from the type locality in the Amatola Mountainsnear Hogsback Eastern Cape Province

Habitat

No further information is available beyond the collection localitywhich is within the Amatola subtropical moist broadleaf forest

Etymology

Amatola a noun in apposition after its collection locality theAmatola forest

Fig5 Parapurcelliaamatola sp nov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotype male in dorsal ventral and lateral views (FndashQ) SEMs of male paratype (F) ventral gonostome complex(G) spiracle (H) ozophore (I) anal region arrowhead indicates opening of the anal gland (J) chelicer (K) chelicer dentition (L) palp arrowhead indicates ventralprocess on trochanter (M) magnified view of anal region (N) tibia and metatarsus of leg II (O) leg II claw (P) leg IV tarsus (Q) magnified view of leg IV tarsusshowing adenostyle Scale bars A B 1mm CndashE 1mm F 01mm G 005mm H 005mm I 01mm J 02mm K 01mm L 02mmM 005mm N 02mmO 005mm P 02mm Q 005mm

Systematic revision of South African pettalids Invertebrate Systematics 385

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 16: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

C

A B

D E

H

I

F G H

I J K L

M

J K L

M N O PN O P

386 Invertebrate Systematics B L de Bivort and G Giribet

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 17: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Parapurcellia convexa sp nov

(Fig 6 Table 3)

Material examined

Holotype lt (NMSA) from Podocarpus forest litter from Weza StateForest near Staffordrsquos Post (30310S 29450E) 30 km from E KokstadKwaZulu Natal Province South Africa leg P M Croeser W Starega26v1985

Paratypes 3 lt 1 (NMSA) same collecting data as holotype

Diagnosis

Dorsal scutum robustly convex higher in the opisthosoma thanthe dorsal width across the ozophores Tergite VIII deeply lobedScopulae of anal plate visible in dorsal view Single opening ofthe anal gland on tergite IX Tergites IVndashVI concave Cheliceraetightly articulated

Description

Male

Total length 173mm width across ozophores 076mmgreatest width 102mm in the opisthosomal area greatestheight 082mm length-width ratio 170 Body light reddish-brown (when preserved in 70 ethanol) Anterior margin ofdorsal scutum concave without lateral projections prosomaroundly triangular Eyes absent Ozophores conical dorsalfacing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 6E K)Transverse prosomal sulcus present but very inconspicuous(Fig 6C) Transverse opisthosomal sulci distinct particularlyin the posterior Mid-dorsal longitudinal opisthosomal sulcusabsent Dorsal scutum convex flattened medially but otherwiseovoid maximum width and height at tergites III and IVrespectively

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 6A D I) near each otherAnterior margin of coxae II endites enlarged curving laterallyand posteriorly to form an m-shaped ridge Endites of coxae IIand III running along their sutures for a short distance givingcoxae III endites a v-shaped appearance coxae IV enditesrunning parallel to coxae IV midline suture for a distancelonger than the gonostome Coxal endites forming smoothsternal plate with greatest width 022mm between coxae IIIand IV Pores of coxal glands visible at inner margins betweencoxae III and IV Sternum absent Coxae IV endites with horn-like projections on anterior margin of gonostome Gonostomeelliptical 0065mm long and 0096mm wide with straightposterior margin and delimited by coxae IV everywhereexcept posterior margin Lateral walls formed by slightlyelevated endites of coxae IV (Fig 6I)

Spiracles nearly circular open laterally with maximumdiameter 0066mm (Fig 6J) Sternal opisthosomal glandsabsent Sternite 8 rectangular Sternites 8 and 9 and tergite IXfree not forming acorona analis (Fig 6L) Tergite IXwith a singlecentral opening of the anal gland oriented posteriorly tergite VIIIlacking anal glands Anal plate measuring 011 025mmdeeply bilobed by an anterior-pointing u-shaped depressionbearing scopulae along the margin of the depression from theposterior margin of the anal plate to the central ventral surface(Fig 6P) Tergites III throughVIII curving anteriorly through themidlinewith tergiteVIII deeply so appearingbilobedTergite IXand anal plate visible in the dorsal view lateral margins of tergiteVII visible in the ventral view Tergite VIII with setae alongits posterior margin Cuticle with tubercular-microgranulatemorphology in all ventral areas including coxae and anal plateexcept on coxal and palpal endites margin of ventral surfacebetween the sternites and tergites and anterior portions ofdistended tergites (Fig 6CndashE IndashL) Granules more widelyspaced than in most other members of Parapurcellia typicallyseparated by distances comparable to the widths of individualgranules

Chelicerae robust with few setae Proximal article withmicrogranulation on dorsal and lateral surfaces with a dorsalprocess but lacking ventral processes (Fig 6B) Second articlesub-cylindrical its widest portion closer to articulation withmobile digit than proximal margin appearing withoutconspicuous ornamentation under light microscopy andbearing a longitudinal apodeme on the lateral side Proximalarticle 055mm long 026mm wide second article 073mmlong 020mm wide moveable finger 025mm long 0055mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each (Fig 6M)

Palp with club-shaped trochanter widest in the anterior(Fig 6B N) Dimensions of palpal articles of male given inTable 3 Palpal claw 0045mm long

Legs (Fig 6B O Table 3) with all claws smooth longand hook-like without lateral ornamentation Surfaces of alltrochanters femurs patellae and tibae entirely ornamentedmetatarsi partially ornamented All tarsi appearing withornamentation on proximal dorsal surfaces and smoothelsewhere by light microscopy Tarsus of leg I lacking distinctsolea Tarsus IV bisegmented (Fig 6B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0050mm bearing no setaedistal margin at 45 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters (Fig 6FndashH) Femaleparatype slightly larger and longer than male holotype Totallength 178mm width across ozophores 075mm greatest width102mm in the opisthosoma nearly as wide (101mm) in theprosoma behind the ozophores greatest height 075m length-

Fig6 Parapurcellia convexa spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsof chelicer palpand legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region (M) chelicer dentition (N) palp(O) tibia and metatarsus of leg II (P) magnified view of anal region arrowhead indicates opening of the anal gland Scale bars A B 1mmCndashH 1mm I 01mmJ 005mm K 01mm L 01mm M 005mm N 02mm O 02mm P 005mm

Systematic revision of South African pettalids Invertebrate Systematics 387

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 18: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

width ratio 175 Anal plate in a terminal position not visible inthe dorsal view without a bilobed elevated anterior margin Notergites bilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality Weza State Forest KwaZuluNatal Province South Africa

Habitat

Specimens were collected in Podocarpus litter

Etymology

From Latin convexus the species is named for the shape of itsopisthosoma in the lateral view which is considerably moreconvex than in other members of this genus

Parapurcellia minuta sp nov

(Fig 7 Table 3)

Material examined

Holotype lt (NMSA) Original label is difficult to read marked asfollows lsquo2820 ndashDurbanAug lsquo40WGRrsquoRecords at theNMSA indicate thefollowing collectednearDurbanKwaZuluNatal Province SouthAfrica legWG Rump viii1940

Paratypes 1 lt 2 (NMSA) same collecting data as holotype

Diagnosis

Small animal TergiteVII concave Chelicerae loosely articulatedwith the anterior prosomal margin Scutum lengthndashwidth rationearly exactly 20 Sternal plate formed by the coxae III and IVendite narrow (016mm)

Description

Male

Total length 152mm width across ozophores 062mmgreatest width 076mm in the prosoma behind the ozophoresthe opisthosoma nearly as wide greatest height 061mm length-width ratio 199 Body pale yellowish-brown (when preservedin 70 ethanol) Anterior margin of dorsal scutum concavewithout lateral projections prosoma roundly triangular Eyesabsent Ozophores conical dorsal facing 45 with terminalozopore with circular opening ornamentation uniform andnon-directional (Fig 7A C E) Transverse prosomal sulcuspresent but inconspicuous (Fig 7C) Transverse opisthosomalsulci inconspicuous Mid-dorsal longitudinal opisthosomalsulcus absent Dorsal scutum flat maximum width and heightin prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 7A D I) near each otherAnteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming small smooth sternal plate with greatest width016mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome elliptical 0056mm long and0081mm wide delimited by coxae IV everywhere exceptposterior margin Lateral walls formed by slightly elevatedendites of coxae IV (Fig 7I)

Spiracles nearly circular open laterally with maximumdiameter 0053mm (Fig 7J) Sternal opisthosomal glandsabsent Posterior margin of sternite 8 curved anteriorly throughthemidline sternite 7 trapezoidal Sternites 8 and 9 and tergite IXfree not forming a corona analis (Fig 7A D L) Tergite IXcurving to the anterior at the mid-line appearing w-shapedTergite IX with a single central opening of the anal glandoriented posteriorly tergite VIII lacking opening of the analglands Anal plate measuring 0072 018mm with ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface along themargins of the raised lateralportions of the anal plate (Fig 7L) Cuticle with tubercular-microgranulate morphology in all ventral areas includingcoxae and anal plate except on coxal and palpal endites andmargin of ventral surface between the sternites and tergites(Fig 7CndashE IndashL P)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process lackingventral processes (Fig 7B) Second article sub-cylindrical itswidest portion nearmiddle of article but closer to articulationwithmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 043mm long 021mm wide second article057mm long 012mm wide moveable finger 021mm long0037mmwide Dentition uniform and similar on both cheliceralfingers with 10 denticles on each (Fig 7M)

Palp with club-shaped trochanter narrowing posteriorlybearing a small ventral process (Fig 7B N) Dimensions ofpalpal articles from trochanter to tarsus of male given inTable 3 Palpal claw 0032mm long

Legs (Fig 7B O Table 3) with all claws smooth long andhook-likewithout lateral ornamentation (Fig 7O) Surfaces of alltrochanters femurs patellae and tibae entirely ornamented

Fig 7 Parapurcelliaminuta sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I II III andIV (CndashE) Automontage photographs of holotypemale in dorsal ventral and lateral views (FndashH) Automontage photographs of paratype female in dorsal ventraland lateral views (IndashP) SEMs of male paratype (I) ventral gonostome complex (J) spiracle (K) ozophore (L) anal region arrowhead indicates opening of theanal gland (M) chelicer dentition (N) palp arrowhead indicates ventral process on trochanter (O) tibia andmetatarsus of leg II (P)magnified viewof anal regionarrowhead indicates opening of the anal gland Scale barsAB 1mmCndashH 1mm I 01mm J 002mmK 005mm L 01mmM 005mmN 02mmO 02mmP 002mm

388 Invertebrate Systematics B L de Bivort and G Giribet

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

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elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

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yi

Sir

o a

caro

ide

s

Tro

glo

siro

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qu

a

Pa

rap

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elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

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oe

dip

us

Ra

kaia

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lita

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ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

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Fa

ng

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sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 19: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

C

II J K L

M N O PN O P

J K L

M

A B

D E

F G H

Systematic revision of South African pettalids Invertebrate Systematics 389

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

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erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

Ra

kaia

so

lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 20: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

metatarsi partially ornamented All tarsi appearing smooth bylight microscopy tarsi of leg I ornamented by a few sparse browngranules Tarsus of leg I lackingdistinct solea (Fig 7B) Tarsus IVbisegmented carrying a small adenostyle towards the middleof the proximal segment terminating in a tuft of ~5 setaeapproximate length of adenostyle 0034mm distal margin at47 of tarsal length

Spermatopositor not studied

Female

Similar to male in non-sexual characters Female paratypeslightly larger and longer than male holotype Total length164mm width across ozophores 060mm greatest width080mm in the opisthosoma nearly as wide (079mm) in theprosoma behind the ozophores greatest height 061mm length-width ratio 206 Anal plate in a similar position as the malewithout a bilobed elevated anterior margin Tergite VIII notbilobed Tarsus IV thin longer than in males notbisegmented Gonostome area typical of pettalids with coxaeof leg IV not meeting in the midline

Distribution

Only known from the type locality which is itself not preciselyknown but is recorded as near Durban KwaZuluNatal ProvinceSouth Africa

Habitat

No further information is available beyond the collection localitywhich has a mild subtropical climate

Etymology

From Latin minutus meaning lsquolessenedrsquo past participle ofminuere the species is named for its small size with a bodymeasuring only 15mm in length

Parapurcellia natalia sp nov

(Fig 8 Table 3)

Material examined

Holotype lt (NMSA) Original label is incomplete marked asfollows lsquo2841 ndash Mazongwa Fa Nov 40 RFLrsquo Records at theNMSA indicate the following collected near Krantzkop MazongwanaForest 20 miles NE of Greytown KwaZulu Natal Province SouthAfrica leg RF Lawrence xi1940

Diagnosis

Small animal widest in prosoma Bearing inconspicuousmid-dorsal longitudinal opisthosomal sulcus Cheliceraetightly articulated with anterior prosomal margin

Description

Male

Total length 158mm width across ozophores 068mmgreatest width 088mm in the prosoma behind the ozophoresgreatest height 057mm length-width ratio 179 Body paleyellowish-brown legs off-white (when preserved in 70ethanol) Anterior margin of dorsal scutum concave without

lateral projections prosoma roundly triangular Eyes absentOzophores conical dorsal facing 45 with terminal ozoporewith circular opening ornamentation uniform and non-directional (Fig 8A C E H) Transverse prosomal sulcuspresent but inconspicuous (Fig 8A C) Transverse opisthosomalsulci visible Mid-dorsal longitudinal opisthosomal sulcusinconspicuous (Fig 8C) Dorsal scutum flat maximum widthand height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 8A D F) Endites ofcoxae II and III running along their sutures giving coxae IIIendites a v-shaped appearance coxae IV endites running parallelto coxae IV midline suture for a distance slightly less thanthe gonostome Coxal endites forming smooth sternal platewith greatest width 028mm between coxae III and IV Poresof coxal glands visible at inner margins between coxae III andIV Sternum absent Coxae IV endites with horn-like projectionson anterior margin of gonostome Gonostome nearly elliptical0080mm long and 0093mm wide and delimited by coxae IVeverywhere except posterior margin Lateral walls formed byelevated endites of coxae IV (Fig 8F)

Spiracles nearly circular open on the posterior half of itslateral side withmaximumdiameter 0053mm (Fig 8G) Sternalopisthosomal glands absent Posterior margin of sternite 8 andsternite 9 curved anteriorly through the midline sternite 7trapezoidal Sternites 8 and 9 and tergite IX free not forming acorona analis (Fig 8D I) Anteriormargin of tergite IX curving tothe anterior at the mid-line appearing deeply bilobed Tergite IXwith a single central opening of the anal gland orientedposteriorly surrounded by a region of small setae Tergite VIIIlacking anal glands Anal plate measuring 012 022mmwith an anterior-pointing v-shaped depression and scopulaeoriginating in the central ventral surface (Fig 8I) Cuticle withtubercular-microgranulate morphology in all ventral areasincluding coxae and anal plate except on coxal and palpalendites margin of ventral surface between the sternites andtergites and depression of anal plate (Fig 8CndashI)

Chelicerae robust with few setae first article withmicrogranulation on lateral and proximal-dorsal surfacessecond article appearing without conspicuous ornamentationunder light microscopy Proximal article with a dorsal processand weak ventral process most prominent laterally (Fig 8B)articulating tightly with anterior margin of prosoma Secondarticle sub-cylindrical its widest portion near articulation withmobile digit bearing a longitudinal apodeme on the lateral sideProximal article 047mm long 021mm wide second article068mm long 015mm wide moveable finger 019mm long0039mmwide Dentition uniform and similar on both cheliceralfingers with 11 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 8B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0036mm long

Legs (Fig 8B Table 3) with all claws appearing smoothlong and hook-like without lateral ornamentation under lightmicroscopy Surfaces of all leg trochanters femurs patellaetibae and metatarsi entirely ornamented All tarsi appearing

390 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

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kaia

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lita

ria

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ita

ca v

en

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lis

Ch

ileo

gov

ea

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sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

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bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 21: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

C

F G H I

A B

D E

CI

CII

CIII

CIV

S1

G

ECII

ECIII

ECIVCG

EP

APsc

opTIX

S9

S8

S7

S6

TVIII

ECI

Fig 8 Parapurcellia natalia sp nov (A) Illustrations of holotype body in dorsal ventral and lateral views (B) Illustrations of chelicer palp and legs I IIIII and IV (CndashE)Automontage photographs of holotype in dorsal ventral and lateral views (FndashI)Automontage photographs of various holotype bodypartswith annotated tracings (F) ventral gonostomecomplexCIndashCIVandEC1ndashECIV indicate the coxae andcoxal enditesof legs IndashIV respectivelyEP is thepalpalendite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plate SC scopulaeon anal plate TIX tergite IXTVIII tergiteVIII andOPopeningof the anal glandDashed line indicates posteriormarginof raised anal plate lobes Scale barsA B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 391

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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Fig12

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ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 22: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

smooth by lightmicroscopy Tarsus of leg I lacking distinct soleaTarsus IV bisegmented (Fig 8B) carrying a lamelliformadenostyle towards the middle of the proximal segmentapproximate length of adenostyle 0069mm bearing no setaedistal margin at 39 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

Distribution

Only known from the type locality 20 miles NE of GreytownKwaZulu Natal Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Natalia a Latinized adjective in the female gender refers to theNatal region of South Africa where the species was collected

Parapurcellia staregai sp nov

(Fig 9 Table 3)

Material examined

Holotype lt (NMSA) Original label is largely illegible marked asfollows lsquo8452 ndash Tra[illegible]W IX-64 RFL[illegible]rsquo Records at theNMSA indicate the following collected in or near Trafalgar CapeProvince South Africa leg RF Lawrence ix1964

Diagnosis

Small animal Chelicerae loosely articulated with anteriorprosomal margin Tergite VIII not deeply convex anal platenot visible in dorsal view Dorsal scutum length-width ratioapproximately 18 Sternite 9 not appearing bilobed

Description

Male

Total length 172mm width across ozophores 074mmgreatest width 100mm in prosoma behind ozophoresgreatest height 068mm length-width ratio 171 Body paleyellowish-brown (when preserved in 70 ethanol) Anteriormargin of dorsal scutum concave without lateral projectionsprosoma roundly triangular Eyes absent Ozophores conicaldorsal facing 45 with terminal ozopore with circular openingornamentation uniform and non-directional (Fig 9A C E H)Transverse prosomal sulcus present but inconspicuous(Fig 9C) Transverse opisthosomal sulci inconspicuous Mid-dorsal longitudinal opisthosomal sulcus inconspicuous Dorsalscutum flat maximum width and height in prosoma

Coxae of legs I II and III free Ventral prosomal complexwithleft and right coxae I notmeeting at themidline instead separatedby the palpal endites coxae II III and IV meeting at the midlinemargin between coxae II and III and margin between coxae IIIand IV both reaching the midline (Fig 9AD F) near each other

Anteriormarginof coxae II endites enlarged curving laterally andposteriorly to form anm-shaped ridge Endites of coxae II and IIIrunning along their sutures giving coxae III endites a v-shapedappearance coxae IVendites runningparallel to coxae IVmidlinesuture for a distance slightly longer than the gonostome Coxalendites forming smooth sternal plate with greatest width034mm between coxae III and IV Pores of coxal glandsvisible at inner margins between coxae III and IV Sternumabsent Coxae IV endites with horn-like projections on anteriormargin of gonostome Gonostome semicircular 0081mm longand 012mm wide with straight posterior margin and delimitedby coxae IV everywhere except posterior margin Lateral wallsformed by elevated endites of coxae IV (Fig 9F)

Spiracles nearly circular open laterally with maximumdiameter 0057mm (Fig 9G) Sternal opisthosomal glandsabsent Sternite 8 curved anteriorly through the midlineposterior margin of sternite 7 similarly curved Sternites 8 and9 and tergite IX free not forming a corona analis (Fig 9D I)Anterior margin of tergite IX curving to the anterior at the mid-line appearingw-shapedTergite IXwith a single central openingof the anal gland oriented posteriorly tergiteVIII lackingopeningof the anal glands Anal platemeasuring 011 021mmwith ananterior-pointing v-shaped depression and scopulae originatingin the central ventral surface (Fig 9I) Cuticle with tubercular-microgranulate morphology in all ventral areas including coxaeand anal plate except on coxal and palpal endites margin ofventral surface between the sternites and tergites and depressionof anal plate (Fig CndashI)

Chelicerae relatively robust with few setae first article withmicrogranulation on dorsal and lateral surfaces second articleappearing without conspicuous ornamentation under lightmicroscopy Proximal article with a dorsal process and weakventral process most prominent laterally (Fig 9B) Second articlesub-cylindrical its widest portion near middle of article butcloser to articulation with mobile digit ornamented by smallscale-like projections and lacking apodemes Proximal article047mm long 022mm wide second article 065mm long014mm wide moveable finger 018mm long 0045mmwide Dentition uniform and similar on both cheliceral fingerswith 12 denticles on each

Palp with club-shaped trochanter narrowing posteriorly(Fig 9B) Dimensions of palpal articles from trochanter totarsus of male given in Table 3 Palpal claw 0033mm long

Legs (Fig 9B Table 3) with all claws appearing smooth longand hook-like without lateral ornamentation under lightmicroscopy Surfaces of all trochanters femurs patellae tibaeand metatarsi entirely ornamented Metatarsi of legs I and IIpartially divided by a radial groove at ~55 of metatarsal lengthAll tarsi appearing smooth by light microscopy Tarsus of leg Ilacking distinct solea Tarsus IV bisegmented carrying alamelliform adenostyle towards the middle of the proximalsegment approximate length 0058mm bearing no setae distalmargin at 48 of tarsal length

Spermatopositor not studied as only the holotype wasavailable

Female

Unknown

392 Invertebrate Systematics B L de Bivort and G Giribet

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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Fig12

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398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 23: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

C

F G H I

A B

D E

APsc

op

TIXS9

S8

S7

S6

TVIII

CI

CII

CIII

CIV

S1

G

ECIII

ECIVCG

EP

ECII

ECI

Fig9 Parapurcellia staregai spnov (A) Illustrationsofholotypebody indorsal ventral and lateral views (B) Illustrationsofchelicer palpand legs I IIIII and IV (CndashE) Automontage photographs of holotype in dorsal ventral and lateral views (FndashI) Automontage photographs of various holotype bodyparts with annotated tracings (F) Ventral gonostome complex CIndashCIV and EC1ndashECIV indicate the coxae and coxal endites of legs IndashIV respectively EP isthe palpal endite CG coxal gland G gonostome and S1 sternite 1 (G) Spiracle (H) Ozophore (I) Anal region S6ndash9 indicate sternites 6ndash9 AP anal plateSC scopulae on anal plate TIX tergite IX TVIII tergite VIII and OP opening of the anal gland Dashed line indicates posterior margin of raised anal platelobes Scale bars A B 1mm CndashE 1mm FndashI 01mm

Systematic revision of South African pettalids Invertebrate Systematics 393

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

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ri

Ra

kaia

me

dia

Pa

rap

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a r

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pia

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Ra

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lata

Pu

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swo

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urc

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Pu

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Ka

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rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

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oe

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us

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kaia

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lita

ria

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 24: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Distribution

Only known from the type locality in or near Trafalgar EasternCape Province South Africa

Habitat

No further information is available beyond the collection localitywhich has a subtropical moist savannah climate

Etymology

Species is named after colleague arachnologistWojciechStaregawhodevoted an important part of his career to the study ofAfricanOpiliones and who curated and identified the Cyphophthalmicollection of the Natal Museum segregating most of the speciesdescribed in this paper

Results and discussion

The mite harvestmen of South Africa currently belong tothree genera in the family Pettalidae Purcellia Hansen ampSoslashrensen 1904 (north-eastern South Africa and CapePeninsula) Speleosiro Lawrence 1931 (Cape Peninsula) andParapurcelliaRosas Costa 1950 (south to eastern SouthAfrica)The phylogenetic analyses below predominantly associatePurcellia peregrinator with Parapurcellia rather than otherspecies in its current genus A suite of morphologicalcharacters unequivocally justifies the transfer of Purcelliaperegrinator to the genus Parapurcellia and forms the basisof rediagnoses of the three SouthAfrican pettalid genera providedabove

In our phylogenetic analyses Fangensis spelaeuswas chosenas the outgroup during heuristic searches but treeswere re-rootedto have the family Stylocellidae as the sister group of theremaining Cyphophthalmi This relationship was found inearlier molecular studies (Giribet and Boyer 2002) particularlywhen using maximum likelihood as optimality criterion (Boyeret al 2007b) These studies also offer support under otherparameters to the hypothesis that the Pettalidae are sister tothe remaining Cyphophthalmi as shown in a recent moreinclusive analysis of the order Opiliones (Giribet et al 2010)The alternative rooting has no effect on the internal phylogenyof Pettalidae

Equally weighted analysis of each data partition

We analysed the discrete and continuous morphology datasetsusing parsimony as the optimality criterion under equal characterweighting Generally fewer than 50 random addition sequenceswere required to find the shortest trees with the discrete datasetFor the discrete data the number of most parsimonious trees islarge (in the thousands) sowe decided to use a driven search untilhittingminimum tree length 1000 times (see Giribet 2005 2007)This yielded 2757 trees of 228 steps which after TBR collapsingresulted in 295 trees The strict consensus of these trees (Fig 10A)has all the families of Cyphophthalmi monophyletic except forSironidae which is unresolved with respect to Pettalidae and(Troglosironidae +Neogoveidae) The monophyly of thesefamilies and the polyphyly of Sironidae are results consistentwith several previous molecular and discrete morphological

studies (Giribet and Boyer 2002 Giribet 2003 de Bivort andGiribet 2004 Boyer et al 2007b Giribet et al 2010)

Optimisation of the characters onto this tree revealsmany character state changes supporting deep family-levelrelationships (eg 11 state changes separate Stylocellidae fromthe other families 16 optimise at the base of Pettalidae and 7 atthe base of (Troglosironidae +Neogoveidae)) However mostcharacter changes are ambiguously optimised changing alongmultiple nodes on the tree Notable unambiguous characterstate changes include the presence of Ramblarsquos organ inFangensis (Schwendinger and Giribet 2005) the presence of asternum in Stylocellidae the presence of a oral lappet in(Troglosironidae +Neogoveidae) the presence of free sternites8 and 9 and tergite IX in Pettalidae the presence of a v-shapedmodification of sternites 6ndash8 in Karripurcellia (Giribet 2003)and the presence of a large anal plate depression in males unitingPurcellia and Speleosiro

Several genera within Pettalidae appear monophyletic inour analyses under equal weighting (Karripurcellia PettalusChileogovea Austropurcellia and Parapurcellia) but theirposition with respect to one another is unstable or receives lowsupport with the exception of Pettalus+Chileogovea and aSouth African clade The New Zealand genera Aoraki andRakaia each monophyletic with high support in all publishedmolecular studies (Boyer and Giribet 2007 2009 Giribet et al2010) are not monophyletic with this dataset In the discrete-character analysis of Boyer and Giribet (2007) both generaare monophyletic but they are supported by a singleambiguous character change and the genera are not recoveredin the continuous-character analysis of de Bivort et al(2010) Speleosiro appears nested within the genus Purcelliaforming a trichotomywith Purcellia illustrans and the remainingPurcellia which form an unresolved clade of animals primarilyfrom western South Africa Parapurcellia is monophyletic withP silvicola and P monticola basal to the remaining species andP amatola and P minuta are sister species The South Africanpettalids form an unresolved clade with Austropurcellia anassociation found in previous molecular and morphologicalstudies (Giribet and Boyer 2002 Boyer et al 2007b Boyerand Giribet 2009 Giribet et al 2010) although these studiessometimes find Purcellia and Austropurcellia to be basal to therest of Pettalidae

Using the continuous morphological character dataset wefound one tree of length 18785 (Fig 10B) that predominantlyagrees with the discrete morphological tree The data in thischaracter matrix underwent independence analysis andcollapse of non-independent characters before analysis usingan independence cutoff value of 131 which was previouslyfound (de Bivort et al 2010) to recover the most family-levelgroups with strong support Thus while it was not surprising thatthe continuous morphological dataset supported all the family-level relationships found in the discrete morphological tree itshould be noted that a broad range of independence cutoff valuessupport those clades (de Bivort et al 2010) In contrast to thediscrete analysis the continuous analysis found Sironidae asmonophyletic with Suzukielus sister to all the other speciesBecause the family-level relationships found in the discretetree are reiterated in the continuous tree and continuous treesby their very nature tend to be fully resolved the strict consensus

394 Invertebrate Systematics B L de Bivort and G Giribet

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

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Systematic revision of South African pettalids Invertebrate Systematics 397

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398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 25: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

of the single continuous tree and thediscrete trees (Fig 10C) looksquite similar to the consensus discrete tree

The concept of optimised character states changing (or not) onparticular branches of a tree does not carry over especially wellfrom discrete to continuous analyses since the degree to which acharacter changes on a branch can vary continuously Thereforewe chose to annotate the continuous character tree with lsquolargersquocharacter changes (Fig 10B) defined (arbitrarily) as at least achange of 15 times the standard deviation of the value of thecharacter across taxaUnlike the changes in the discrete tree deepfamily-level branches of the tree were not associated with largechanges in values of the continuous characters Large changesto characters that occurred only once on the tree include thewidening of the spiracle opening in Stylocellidae the wideningof bilateral features of the scutum (such as the ozophores andthe terminal posterior lobes) in Pettalidae and the increasedconcavity of tergite VI in Aoraki longitarsa+Pettalus

The internal relationships of the pettalid genera supportedby the continuous tree differ from those few intergenericrelationships supported by the discrete tree For exampleChileogovea sister to Pettalus in the discrete tree is foundalong with Karripurcellia in a grade at the base of PettalidaeAndwhileAoraki andRakaiawereunresolved in the discrete treein the continuous tree they cluster in a clade that also includesPettalus (from Sri Lanka) and Speleosiro (from South Africa)a relationship that is unlikely given the geologic history ofGondwana the low vagility in these animals molecularevidence in favour of the monophyly of each of the NewZealand pettalid genera (Boyer and Giribet 2007) andmorphological evidence for the monophyly of Purcellia plusSpeleosiro (Giribet 2003) Nevertheless the continuous treeagrees with the discrete tree about several relationshipswithin Pettalidae including the monophyly of Parapurcelliathe association of Parapurcellia monticola and P silvicola

Fig 10 (A) Strict consensus of thousands of trees each of length 228 obtainedunder parsimony analysis of the discrete character dataset under equalweightingLabels on the right indicate families Hollow boxes indicate changes in ambiguous characters ndash defined as characters that changemore than once in the tree Filledboxes indicate unambiguous characters Small numbers are the number of the character changing Italicised numbers on branches indicate the bootstrap supportvalue (B) Shortest tree (length 18785) obtained under parsimony analysis of the continuous character dataset Since most continuous character change to somedegree on all branchesmarks indicate those character changes ofmagnitudegreater than15 standarddeviations of the characterrsquos values across all taxaMarkingsare otherwise the same as on the discrete tree (C) Strict consensus of trees A and B

Systematic revision of South African pettalids Invertebrate Systematics 395

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

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ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

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ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

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Ka

rrip

urc

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oru

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arr

ipu

rce

llia

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rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

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oif

fait

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Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

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a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

vea

oe

dip

us

Ra

kaia

so

lita

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Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

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dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

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nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

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urc

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a s

tare

ga

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nov

Pe

tta

lus

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rmis

Pa

rap

urc

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a f

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Fa

ng

en

sis

spe

lae

us

Tro

glo

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ae

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Pa

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a s

ilvic

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Sty

loce

llus

tam

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Pa

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urc

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on

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la

Pa

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urc

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a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

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wo

od

wa

rdi

Tro

glo

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lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

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en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 26: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

P minuta and P amatola as sister species and the positioning ofthe South African pettalids as derived within the family ndash arelationship typically supported by morphological (Giribet andBoyer 2002) but not molecular data (Boyer et al 2007b)

Implied weighting analysis of each data partition

Analysis of either the discrete or morphological datasets underimplied weighting using a variety of concavity values (k) yieldedtrees with a variety of topologies Therefore the consensus ofimplied weighting trees which reveals groups with robustsupport in the dataset for each dataset (Fig 11) was largelyunresolved particularly the consensus tree generated fromcontinuous morphological data only

The consensus of all most parsimonious trees derivedfrom the discrete dataset (Fig 11A) with k ranging from 1 to 6(trees for k gt 6 were essentially identical to the equally weightedtree) had all families as monophyletic with the exception ofSironidae which was polyphyletic by the placement ofSuzukielus as the sister group of Pettalidae This relationshipis not unprecedented in discrete datasets (de Bivort and Giribet2004) and the position of Suzukielus varies between treeswith different optimisation criteria in analyses of combinedmolecular and discrete datasets (Boyer et al 2007b) Therelationships between genera within Pettalidae were nearlyentirely unresolved within this tree although Speleosiro wasfound as sister to Purcellia illustrans and Purcellia was sisterto Parapurcellia peregrinator This latter relationship is notablebecause this species was originally described in the genusPurcellia mdash as were all the South African species describedby RF Lawrence even after JA Rosas Costa erected the genusParapurcellia in 1950)mdash andwas re-assigned above Among allthe trees presented herein only this one supports Parapurcelliaperegrinator as more closely allied with Purcellia thanParapurcellia

For values of k ranging from 3 to 6 the consensus of treesderived from the discrete dataset (Fig 11B) is considerably moreresolved providing some support forChileogovea as sister to therest of Pettalidae with Pettalus +Karripurcellia as the next mostbasal group Austropurcellia was found as the sister group of theSouthAfrican pettalids inwhichPurcellia is found nestedwithinParapurcellia

Under implied weighting with k ranging from 1 to 6 thecontinuous dataset yielded trees whose strict consensus(Fig 11C) is nearly entirely unresolved supporting onlyStylocellidae Austropurcellia Parapurcellia amatola+P natalia +P minuta and the highly improbable groupTroglosironidae + Siro acaroides+Cyphophthalmus duricoriusThe consensus of trees with k ranging only from 4 to 6(Fig 11D) reveals support for a few plausible groups such asPurcellia AustropurcelliaKarripurcellia andPettalus Becausethe continuous dataset was prepared using independence analysisand collapse of dependent characters all the characters usedin the analysis contain unique phylogenetic information ndash iethere is minimal redundancy between characters Perhaps thelow quality of the trees generated with continuous data andimplied weighting is due to down-weighting of non-redundantphylogenetic information by the implied weighting algorithmIndeed the continuous dataset was highly sensitive to implied

weighting and only yielded trees similar to those from the equallyweighted analysis for k gt 30

Analysis of combined data under equaland implied weighting

The most parsimonious trees generated using either discreteor continuous data under equal weighting and discrete dataunder implied weighting support numerous family-levelrelationships (Figs 10 11) Moreover both continuous anddiscrete datasets under equal weighting support severalspecific relationships within Pettalidae (such as Parapurcelliaamatola and P minuta being sister species) (Fig 10) Becauseboth datasets support several evolutionary relationships weperformed phylogenetic analyses on datasets combining thesetwo data partitions under weightings designed to vary theirrespective contributions to the final tree length (see Methodssection) These analyses were done by allowing the characters tobe differentially weighted in two different ways simultaneouslyFirst the discrete and continuous data partitions were weightedrelative to each other (in terms of their approximate finalrespective contributions to tree length ndash see Methods) asfollows 8 1 4 1 2 1 1 1 1 2 1 4 and 1 8 Second foreachof thosepartitionweightingsTNTanalyseswere doneunderequal weighting and implied weighting for k= 1ndash6 This yielded49 different trees The status of many groups of interest acrossthese parameter conditions is shown in Fig 12A The supportedphylogenetic groups tend to be robust to the relative weighting ofthe continuous and discrete data partitions but sensitive to theconcavity parameter of the implied weighting analysis (k)

Many plausible groups are found to be monophyleticunder all values of k and all relative weightings between thecontinuous and discrete data partitions ndash the families PettalidaeStylocellidae Neogoveidae and Troglosironidae the cladeNeogoveidae + Troglosironidae the genera KarripurcelliaAustropurcellia Chileogovea and Pettalus and the sisterrelationship of Purcellia griswoldi and P lawrencei Withrespect to Pettalidae Suzukielus is found as its sister group inall implied weighting analyses consistent with the impliedweighting analysis of the discrete data alone (Fig 11A B) Inall equal weighting analyses Suzukielus is found withinSironidae The position of Suzukielus has been contentious Inprevious morphological analyses of discrete data Suzukieluswasallied to Pettalidae (Giribet and Boyer 2002 Giribet 2003de Bivort and Giribet 2004) but it was placed with Sironidaein the analysis of continuous character data of de Bivort et al(2010) In all analyses here presented there is an affinity betweenPettalidae and Sironidae with the two families forming a grade inmost analyses that includesTroglosironidae +Neogoveidae) anda clade for high k values and datasets in which the discretepartition is weighted highly

Within Pettalidae the genera Aoraki and Rakaia which arerespectively monophyletic in molecular studies (Boyer andGiribet 2007 2009 Boyer et al 2007b) were not found to bemonophyletic although in most analyses in which k is not equalto 1 the two genera form a grade Karripurcellia and Pettalusare sister genera in 27 of 49 cases with little relation to k orthe partition weighting The South African genus Parapurcelliawas always found as either a non-monophyletic grade at the

396 Invertebrate Systematics B L de Bivort and G Giribet

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

illu

stra

ns

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

go

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oe

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us

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kaia

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lita

ria

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ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

llus

ram

bla

e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 27: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

AB

CD

Fig11

(A)S

trictcon

sensus

oftreesob

tained

underparsimon

yanalysisofthediscretedatasetund

erim

pliedweightsfork

valuesrang

ingfrom

1to6(B)A

sinAbuton

lyfork

rangingfrom

3to6(C)S

trict

consensusof

treesobtained

underparsim

onyanalysisof

thecontinuous

datasetunderim

pliedweightsforkvalues

rang

ingfrom

1to

6(D

)Asin

Cb

uton

lyforkrangingfrom

4to

6

Systematic revision of South African pettalids Invertebrate Systematics 397

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

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swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

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elli

a p

eck

oru

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arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

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es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

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wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

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qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

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stra

ns

Cyp

ho

ph

tha

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s d

uri

cori

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Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

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oe

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us

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en

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Ch

ileo

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sta

Fa

ng

en

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cave

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Fa

ng

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loce

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sisi

Pa

rasi

ro c

oif

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i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

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lam

pe

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es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

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ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

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elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

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ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 28: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

EW

12

3

IW c

onca

vity

(k)

poly

phyl

etic

para

phyl

etic

mon

ophy

letic

approx tree length contributionratio (continuousdiscrete)

45

6

81

41

21

11

12

14

18

Pet

talid

ae S

tylo

celli

dae

Neo

gove

idae

Tro

glos

ironi

dae

(T

rogl

osiro

nida

e amp

Neo

gove

idae

)K

arrip

urce

llia

Aus

trop

urce

llia

C

hile

ogov

ea P

etta

lus

(Pur

celli

a gr

isw

oldi

sp

nov

ampP

urce

llia

law

renc

ei s

p n

ov)

(Pet

talid

ae +

Suz

ukie

lus)

(Aor

aki +

Rak

aia)

(Pet

talid

ae +

Siro

nida

e)

(Kar

ripur

celli

a +

Pet

talu

s)

(Par

apur

celli

a)

(Pur

celli

a +

Par

apur

celli

a+

Spe

leos

iro)

(Pur

celli

a +

Par

apur

celli

a +

Spe

leos

iro +

Aus

trop

urce

llia)

AB

C

Ao

raki

cry

pta

Ao

raki

de

nti

cula

Su

zuki

elu

s sa

ute

ri

Ra

kaia

me

dia

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

iso

lata

Pu

rce

llia

gri

swo

ldi

sp

no

vP

urc

elli

a t

ran

sva

alic

a

Ao

raki

lo

ng

ita

rsa

Pu

rce

llia

law

ren

cei s

p n

ov

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

mK

arr

ipu

rce

llia

ha

rve

yi

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Ra

kaia

so

ren

sen

i dig

itata

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a p

ere

gri

na

tor

Ra

kaia

ma

gn

a a

ust

ralis

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Sir

o r

ub

en

s

Pa

rasi

ro c

oif

fait

i

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Ne

og

ove

a s

p

Pa

rap

urc

elli

a f

issa

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a m

on

tico

la

Pe

tta

lus

lam

pe

tid

es

Ao

raki

tu

mid

ata

Me

tag

ove

a p

hili

pi

Au

stro

pu

rce

llia

wo

od

wa

rdi

Ra

kaia

lin

dsa

yi

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a

Pa

rap

urc

elli

a s

ilvic

ola

Ao

raki

in

erm

a

Pu

rce

llia

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stra

ns

Cyp

ho

ph

tha

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s d

uri

cori

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Sp

ele

osi

ro a

rga

sifo

rmis

Ch

ileo

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oe

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us

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ita

ca v

en

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lis

Ch

ileo

gov

ea

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sta

Fa

ng

en

sis

cave

rna

rum

Sty

loce

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e

Fa

ng

en

sis

spe

lae

us

Sty

loce

llus

tam

bu

sisi

Pa

rasi

ro c

oif

fait

i

Fa

ng

en

sis

cave

rna

rum

Pu

rce

llia

illu

stra

ns

Ao

raki

tu

mid

ata

Pe

tta

lus

lam

pe

tid

es

Me

tag

ove

a p

hili

pi

Su

zuki

elu

s sa

ute

ri

Hu

ita

ca v

en

tra

lis

Ch

ileo

gov

ea

joca

sta

Ne

og

ove

a s

p

Ra

kaia

so

ren

sen

i d

igit

ata

Au

stro

pu

rce

llia

sco

pa

ria

Ka

rrip

urc

elli

a p

eck

oru

m

Pa

rap

urc

elli

a a

ma

tola

sp

nov

Pu

rce

llia

law

ren

cei s

p n

ov

Ch

ileo

go

vea

oe

dip

us

Sty

loce

llus

ram

bla

e

Ka

rrip

urc

elli

a s

ierw

ald

ae

Pu

rce

llia

tra

nsv

aa

lica

Pa

rap

urc

elli

a m

inu

ta s

p n

ov

Pa

rap

urc

elli

a s

tare

ga

i sp

nov

Pe

tta

lus

cim

icifo

rmis

Pa

rap

urc

elli

a f

issa

Fa

ng

en

sis

spe

lae

us

Tro

glo

siro

ae

llen

i

Pa

rap

urc

elli

a s

ilvic

ola

Sty

loce

llus

tam

bu

sisi

Pa

rap

urc

elli

a m

on

tico

la

Pa

rap

urc

elli

a n

ata

lia s

p n

ov

Ao

raki

cry

pta

Ra

kaia

ma

gn

a a

ust

ralis

Pu

rce

llia

gri

swo

ldi

sp

no

v

Sp

ele

osi

ro a

rga

sifo

rmis

Au

stro

pu

rce

llia

wo

od

wa

rdi

Tro

glo

siro

lo

ng

ifoss

a

Pa

rap

urc

elli

a c

onv

exa

sp

nov

Sir

o r

ub

en

s

Pa

rap

urc

elli

a p

ere

gri

na

tor

Cyp

ho

ph

tha

lmu

s d

uri

cori

us

Pa

rap

urc

elli

a r

um

pia

na

Ra

kaia

so

lita

ria

Ra

kaia

iso

lata

Ao

raki

lo

ng

ita

rsa

Ra

kaia

lin

dsa

yi

Ra

kaia

me

dia

Ao

raki

in

erm

a

Ao

raki

de

nti

cula

Sir

o a

caro

ide

s

Tro

glo

siro

nin

qu

a Ka

rrip

urc

elli

a h

arv

eyi

99 51 69

75 92 95 8581

80 69

94

6872

9998

74 83

86 86 5665

6379

8877 100

9381

81 87

68

97

Fig12

(A)Navajorugs

summarisingresults

forcladesof

interestanalysed

underparsim

onyusingcombineddiscreteandcontinuous

dataT

hesearch

conditionsvaried

acrossequalw

eights(EW)and

impliedweightswith

kvalues

rang

ingfrom

1to6andtherelativ

eweigh

tofthe

continuous

datasettothediscretedatasetranging

from

81

to18(B)S

trictconsensus

ofalltrees

foun

dinwhich

thegenus

Parap

urcelliaisfoun

dtobe

aderivedgrou

pwith

inthePettalid

ae(and

mon

ophyletic)Thiscorrespon

dstoalltreesun

derE

Wk=4k=5andk=6andk=3fordatapartition

weightin

gratio

sof8

11

11

2

14

and18(C)S

trictcon

sensus

ofalltreesinwhich

thegenusParapurcelliaisfoun

dtobe

basalw

ithinthePettalid

ae(and

paraphyletic)Thiscorrespo

ndstoalltreesun

derk

=1andk=2andk=3ford

ata

partition

weightin

gratio

sof41

and21N

umberson

thebranchesoftreesinBandCindicatethebootstrapsupportvaluesderivedfrom

analysesofthe11

weightedcombineddatasetunderEWk

=1and

k=6respectiv

ely

398 Invertebrate Systematics B L de Bivort and G Giribet

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 29: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

base of Pettalidae (for k= 1 2 and sometimes 3) or a cladeThis distinction can be used to establish two alternativehypotheses for the phylogenetic relationships within Pettalidae(Fig 12B C)

The strict consensus of all trees in which Parapurcellia wasfound to be monophyletic (Fig 12B) places the South Africanpettalids in a derived internally unresolved clade that alsoincludes Austropurcellia The affinity of Austropurcellia andthe South African pettalids was also found in the molecularstudy Boyer et al (2007b) although they also found aconsistent association between the South African genera andthe South American genus Chileogovea that was absent in ourresults The strict consensus of all remaining trees (Fig 12C) isconsistent with Boyer et al (2007b) in that Parapurcelliaforms a grade at the base of Pettalidae In our tree Purcelliaand Speleosiro form a clade which is in turn found in anunresolved group containing the remaining pettalid generaBoth of these evolutionary hypotheses have respectiveconsistencies with previous molecular studies thus it is hardfor us to favour one over the other on the basis of phylogeneticrelationships found in earlier studies

We performed bootstrap analysis on the dataset with 1 1weighting between the partitions for k= 1 and k= 6 whichrespectively correspond to the derived (Fig 12B) and basal(Fig 12C) hypotheses for the position of ParapurcelliaBoth analyses yielded support values gt50 for StylocellidaeTroglosironidae Pettalidae Troglosironidae +Neogoveidaesix pettalid genera Cyphophthalmus+ Siro rubens andPurcellia griswoldi+P lawrencei However (for theseparameters) there is greater support for several other clades inthe basal Parapurcellia hypothesis NeogoveidaeParapurcelliaconvexa+P silvicola Aoraki crypta+A inerma (Boyer et al2007b) Speleosiro +Purcellia Purcellia and P illustrans + theremaining Purcellia This may provide some evidence in favourof the hypothesis that places Parapurcellia basal withinPettalidae

Position of Parapurcellia peregrinator

Our reassignment of Purcellia peregrinator to Parapurcelliaperegrinator is supported by numerous discrete morphologicalcharacter differences between these genera eg the presenceabsence of anterior projections of the gonostome wall theposteriorventral orientation of the anal gland opening and theabsencepresence of a large anal plate depression While therewere no large magnitude changes in the value of continuouscharacters between Purcellia and Parapurcellia this speciesnevertheless falls well within Parapurcellia in the continuousdata analyses

Only in the analysis of the discrete data alone under impliedweighting does Parapurcellia peregrinator occur outsideParapurcellia in this case as the sister group to Purcellia +Speleosiro This does not imply that P peregrinator belongswithin Purcellia since Parapurcellia is rendered paraphyleticwith respect to Purcellia+ Speleosiro in this tree Lastly in theanalysis of the combined discrete and continuous datasetsP peregrinator nests within Parapurcellia whether the genusis found as a derived clade within or at the base of PettalidaeIn the phylogenetic hypothesis in which Parapurcellia is basal

P peregrinator is found as the sister group to the remainingnon-Parapurcellia pettalids (Fig 12C) From these observationswe conclude that Parapurcellia peregrinator is clearly not amember ofPurcellia and instead belongs in the reassigned genusthough it appears in a few instances to be an intermediate taxonbetween the rest of Parapurcellia and either Purcellia or the restof the family Pettalidae

Biogeography of the South African pettalids

In a cladistic biogeographic analysis of the distributions of spidersacross Afromontane regions Griswold (1991) found that theKnysna and Table Mountain areas of endemism are sisterregions while the eastern regions are more closely related toother regions outside of South Africa namely a clade formedby the ((Eastern Africa Volcanoes + Eastern Arc Mountains) +Madagascar) The eastern regions show a sister relationshipbetween the Natal Drakensberg and the Transkei-NatalMidlands while the Transvaal Drakensberg is more closelyrelated to the Africa Volcanoes +Eastern Arc Mountains +Madagascar clade We found that Purcellia and Speleosiroform a clade supporting in part the relationships of theKnysna and Table Mountain areas of endemism butP transvaalica is found in Griswoldrsquos Transvaal Drakensbergarea of endemism which is more closely related to the((Africa Volcanoes +Eastern Arc Mountains) +Madagascar)clade The distribution of Parapurcellia corresponds withGriswoldrsquos Natal-Zululand Coast Transkei-Natal Midlandsand Natal Drakensberg areas of endemism but ourphylogenetic conclusions are too tenuous to derive meaningfulcomparisons

One can compare the distribution of pettalids with thedistributions of other terrestrial invertebrates with likelysimilar dispersal rates and ecological requirements (Harvey1996) Among Opiliones several groups show similardistribution patterns but perhaps most striking is that of thegenera Graemontia Lawrence 1931 as recently summarised byKury (2006) andMonomontiaLawrence 1931 (seeKauri 1961)Phylogenetic analyses of these genera would be necessarybefore concluding whether their biogeographic patterns arealso similar Another matching distribution is that of theonychophoran genus Peripatopsis (Hamer et al 1997) but inthis case it seems that there are several basal lineages in theWestern Cape province (Table Mountain and Knysna Forestareas of endemism) and from there the clade has extended itsrange towards the Eastern Cape and Kwazulu-Natal provinces(Daniels et al 2009)

Note added in proof

During the course of this study a new South African speciesPurcellia leleupi Starega 2008 was described in the PolishJournal of Entomology 77(1) 51ndash56 We were not aware ofthis species until our study was completed Its description andillustrations do not allow clear diagnosis of the species but itseems that the apical tarsal segment lengthndashwidth ratio (~30)distinguishes it fromP griswoldi andP lawrencei and sternite 8looks very different being completely divided and triangular oneach side although this could be an inaccuracy of the description

Systematic revision of South African pettalids Invertebrate Systematics 399

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 30: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Acknowledgements

We are very thankful to A Ndaba (NMSA) for help clarifying old collectiondata from their collection and L Benavides for assistance with lightmicroscopy imaging of the smallest animals The following colleagues arekindly acknowledged for providing specimens for our study N Platnick andL Prendini (AMNH) J Beccaloni (BMNH) S Pollard (CM) P Sierwald(FMNH) P Schwendinger (MHNG) A Muntildeoz-Cuevas (MNHN) P Sirvid(MNZ) R Pinto-da-Rocha (MZUSP) C Conway and A Ndaba (NMSA)D Walter (QM) M Cochrane (SAM) M Grashoff and P Jaumlger (SMF)J Coddington (USNM) M Harvey (WAM) J Dunlop (ZMB) H Dastych(ZMH) and N Scharff (ZMUC) P Schwendinger assisted with speciesnames etymology The Willi Hennig Society sponsored free publiclyavailable versions of TNT Two anonymous reviewers and associate editorNikolaj Scharff provided detailed comments that helped to improve an earlierversion of this article This material is based upon work supported by theNational Science Foundation under Grant no 0236871

References

Boyer S L and Giribet G (2007) A new model Gondwanan taxonsystematics and biogeography of the harvestman family Pettalidae(Arachnida Opiliones Cyphophthalmi) with a taxonomic revision ofgenera from Australia and New Zealand Cladistics 23 337ndash361doi101111j1096-0031200700149x

Boyer S L and Giribet G (2009) Welcome back New Zealand regionalbiogeography and Gondwanan origin of three endemic genera ofmite harvestmen (Arachnida Opiliones Cyphophthalmi) Journal ofBiogeography 36 1084ndash1099 doi101111j1365-2699200902092x

Boyer S L Baker J M and Giribet G (2007a) Deep genetic divergencesin Aoraki denticulata (Arachnida Opiliones Cyphophthalmi) awidespread lsquomite harvestmanrsquo defies DNA taxonomy MolecularEcology 16 4999ndash5016 doi101111j1365-294X200703555x

Boyer S LClouseRMBenavides LR SharmaP Schwendinger P JKarunarathna I and Giribet G (2007b) Biogeography of the worlda case study from cyphophthalmidOpiliones a globally distributed groupof arachnids Journal of Biogeography 34 2070ndash2085 doi101111j1365-2699200701755x

Clouse R M and Giribet G (2007) Across Lydekkerrsquos line-first reportof mite harvestmen (Opiliones Cyphophthalmi Stylocellidae) fromNew Guinea Invertebrate Systematics 21 207ndash228 doi101071IS06046

Clouse R M de Bivort B L and Giribet G (2009) A phylogeneticanalysis for the south-east Asian mite harvestman family Stylocellidae(Opiliones Cyphophthalmi) ndash a combined analysis using morphometricand molecular data Invertebrate Systematics 23 515ndash529 doi101071IS09044

Daniels S R Picker M D Cowlin R M and Hamer M L (2009)Unravelling evolutionary lineages among South African velvet worms(Onychophora Peripatopsis) provides evidence for widespread crypticspeciation Biological Journal of the Linnean Society Linnean Societyof London 97 200ndash216 doi101111j1095-8312200901205x

de Bivort B and Giribet G (2004) A new genus of cyphophthalmidfrom the Iberian peninsula with a phylogenetic analysis of theSironidae (Arachnida Opiliones Cyphophthalmi) and a SEM databaseof external morphology Invertebrate Systematics 18 7ndash52 doi101071IS03029

de Bivort B L Clouse R M and Giribet G (2010) A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen(Opiliones Cyphophthalmi Pettalidae) Journal of ZoologicalSystematics and Evolutionary Research 48 294ndash309 doi101111j1439-0469200900562x

Ewing H E (1923) Holosiro acaroides new genus and species the onlynew world representative of the mite-like phalangids of the suborderCyphophthalmi Annals of the Entomological Society of America 16387ndash391

Forster R R (1948) The sub-order Cyphophthalmi Simon in New ZealandDominion Museum Records in Entomology 1 79ndash119

Forster RR (1952) Supplement to the sub-orderCyphophthalmiDominionMuseum Records in Entomology 1 179ndash211

Forster R R (1955) Further Australian harvestmen (Arachnida Opiliones)Australian Journal of Zoology 3 354ndash411 doi101071ZO9550354

Giribet G (2000) Catalogue of the Cyphophthalmi of the world (ArachnidaOpiliones) Revista Ibeacuterica de Aracnologiacutea 2 49ndash76

Giribet G (2002) Stylocellus ramblae a new stylocellid (OpilionesCyphophthalmi) from Singapore with a discussion of the familyStylocellidae The Journal of Arachnology 30 1ndash9 doi1016360161-8202(2002)030[0001SRANSO]20CO2

Giribet G (2003) Karripurcellia a new pettalid genus (Arachnida Opiliones Cyphophthalmi) from Western Australia with a cladisticanalysis of the family Pettalidae Invertebrate Systematics 17387ndash406 doi101071IS02014

Giribet G (2005) Book reviews TNT tree analysis using new technologySystematic Biology 54 176ndash178 doi10108010635150590905830

Giribet G (2007) Efficient tree searches with available algorithmsEvolutionary Bioinformatics 3 1ndash16

Giribet G and Boyer S L (2002) A cladistic analysis of thecyphophthalmid genera (Opiliones Cyphophthalmi) The Journal ofArachnology 30 110ndash128 doi1016360161-8202(2002)030[0110ACAOTC]20CO2

Giribet G and Boyer S L (2007) Pettalidae Shear 1980 In lsquoHarvestmenThe Biology of Opilionesrsquo (Eds R Pinto-da-Rocha G Machadoand G Giribet) pp 99ndash101 (Harvard University Press CambridgeMA)

Giribet G and Prieto C E (2003) A new AfrotropicalOgovea (OpilionesCyphophthalmi) from Cameroon with a discussion on the taxonomiccharacters in the family Ogoveidae Zootaxa 329 1ndash18

Giribet G Vogt L PeacuterezGonzaacutelez A Sharma P andKury A B (2010)A multilocus approach to harvestman (Arachnida Opiliones) phylogenywith emphasis on biogeography and the systematics of LaniatoresCladistics 26 408ndash437 doi101111j1096-0031200900296x

Goloboff P A (1993) Estimating character weights during tree searchCladistics 9 83ndash91 doi101111j1096-00311993tb00209x

Goloboff P A (1999) Analyzing large data sets in reasonable timessolutions for composite optima Cladistics 15 415ndash428 doi101111j1096-00311999tb00278x

Goloboff P A Farris J S and Nixon K C (2008) TNT a free programfor phylogenetic analyses Cladistics 24 774ndash786 doi101111j1096-0031200800217x

Goodnight C J and Goodnight M L (1980) Metagovea philipi n spa new cyphophthalmid (Arachnida) from Ecuador Transactions of theAmerican Microscopical Society 99 128ndash131 doi1023073226087

Griswold C E (1991) Cladistic biogeography of Afromontane spidersAustralian Systematic Botany 4 73ndash89 doi101071SB9910073

Hamer M L Samways M J and Ruhberg H (1997) A review of theOnychophora of South Africa with discussion of their conservationAnnals of the Natal Museum 38 283ndash312

Hansen H J (1921) lsquoStudies on Arthropoda I The Pedipalpi Ricinuleiand Opiliones (exc Op Laniatores) collected by Mr Leonardo Fea intropical West Africa and adjacent Islandsrsquo (Gyldendalske BoghandelCopenhagen)

Hansen H J and Soslashrensen W (1904) lsquoOn two orders of ArachnidaEspecially the Suborder Cyphophthalmi and Ricinulei Namely theFamily Cryptostemmatoidaersquo (Cambridge University Press CambridgeUK)

Harvey M S (1996) Small arachnids and their value in Gondwananbiogeographic studies In lsquoGondwanan Heritage Past Present andFuture of Western Australian Biotarsquo (Ed S D Hopper) pp 155ndash162(Surrey Beatty amp Sons Sydney)

Hirst S (1925) On some new genera and species of Arachnida Proceedingsof the Zoological Society of London 1925 1271ndash1280

400 Invertebrate Systematics B L de Bivort and G Giribet

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 31: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Joseph G (1868) Cyphophthalmus duricorius eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdecktin der Lueumlger Grotte in Krain Berliner Entomologische Zeitung 12241ndash250 doi101002mmnd18680120208

Juberthie C (1956) Une nouvelle espegravece drsquoOpilions Sironidae de France etdrsquoEspagne Parasiro coiffaiti n sp Bulletin du Museacuteum nationaldrsquohistoire naturelle Paris 28 394ndash400

Juberthie C (1969) Sur les opilions cyphophthalmes Stylocellinae duGabon Biologia Gabonica 5 79ndash92

Juberthie C (1970) Les opilions cyphophthalmes cavernicoles Notes surSpeleosiro argasiformis Lawrence Bulletin du Museum NationaldrsquoHistoire Naturelle 42 864ndash871

Juberthie C (1979) Un cyphophthalme nouveau drsquoune grotte de Nouvelle-Caledonie Troglosiro aelleni n gen n sp (Opilion Sironinae) RevueSuisse de Zoologie 86 221ndash231

Juberthie C (1988) Un nouvel opilion cyphophthalme aveugle drsquoAustralieAustropurcellia gen nov scoparia n sp Memoires de Biospeologie15 133ndash140

Kauri H (1961) Opiliones In lsquoSouth African Animal Life Results of theLundUniversity Expedition in 1950ndash1951rsquo (Eds B Hanstroumlm P Brinckand G Rudebeck) pp 9ndash197 (Almquist amp Wiksell Uppsala)

Kury A B (2006) A new species of Graemontia Lawrence 1931 from theWestern Cape South Africa with notes on the relationships of the genus(Opiliones Laniatores Triaenonychidae) African Zoology 41 45ndash50doi1033771562-7020(2006)41[45ANSOGL]20CO2

Latreille P A (1804) lsquoHistoire naturelle generale et particuliegravere desCrustaceacutes et des Insectesrsquo (F Duart Paris)

Lawrence R F (1931) The harvest-spiders (Opiliones) of South AfricaAnnals of the South African Museum 29 341ndash508

Lawrence R F (1933) The harvest-spiders (Opiliones) of Natal Annalsof the Natal Museum 7 211ndash241

Lawrence R F (1939) A contribution to the opilionid fauna of Natal andZululand Annals of the Natal Museum 9 225ndash243

Lawrence R F (1963) The Opiliones of the Transvaal Annals of theTransvaal Museum 24 275ndash304

Legg G (1990) Parogovia pabsgarnoni sp n (Arachnida OpilionesCyphophthalmi) from Sierra Leone with notes on other Africanspecies of Parogovia Bulletin of the British Arachnological Society8 113ndash121

Murphree C S (1988)Morphology of the dorsal integument of ten opilionidspecies (ArachnidaOpiliones)TheJournalofArachnology16 237ndash252

Nixon K C (1999) The Parsimony Ratchet a new method for rapidparsimony analysis Cladistics 15 407ndash414 doi101111j1096-00311999tb00277x

Pickard-CambridgeO (1875)On three newand curious forms ofArachnidaAnnals and Magazine of Natural History Series 4 383ndash390

Roewer C F (1916) 7 neue Opilioniden des Zoolog Museums in BerlinArchiv fuumlr Naturgeschichte 81 6ndash13

Roewer C F (1923) lsquoDie Weberknechte der Erde SystematischeBearbeitung der bisher bekannten Opilionesrsquo (Verlag von GustavFisher Jena)

Roewer C F (1961) Opiliones aus Sud-Chile Senckenbergiana Biologica42 99ndash105

Rosas Costa J A (1950) Sinopsis de los geacuteneros de Sironinae con ladescripcioacuten de dos geacuteneros y una especie nuevos (OpilionesCyphophthalmi) Arthropoda 1 127ndash151

Schwendinger P J and Giribet G (2005) The systematics of the south-east Asian genus Fangensis Rambla (Opiliones Cyphophthalmi Stylocellidae) Invertebrate Systematics 19 297ndash323 doi101071IS05023

Sharma P and Giribet G (2005) A new Troglosiro species (OpilionesCyphophthalmi Troglosironidae) from New Caledonia Zootaxa 105347ndash60

Sharma P and Giribet G (2006) A new Pettalus species (OpilionesCyphophthalmi Pettalidae) from Sri Lanka with a discussion on theevolution of eyes in Cyphophthalmi The Journal of Arachnology 34331ndash341 doi101636H05-331

Shear W A (1979)Huitaca ventralis n gen n sp with a description of agland complex new to cyphophthalmids (Opiliones Cyphophthalmi)The Journal of Arachnology 7 237ndash242

Shear W A (1985) Marwe coarctata a remarkable new cyphophthalmidfrom a limestone cave in Kenya (Arachnida Opiliones) AmericanMuseum Novitates 2830 1ndash6

Shear W A (1993a) The genus Chileogovea (Opiliones CyphophthalmiPettalidae) The Journal of Arachnology 21 73ndash78

Shear W A (1993b) The genus Troglosiro and the new familyTroglosironidae (Opiliones Cyphophthalmi) The Journal ofArachnology 21 81ndash90

Shear W A (1993c) New species in the opilionid genus Stylocellus fromMalaysia Indonesia and the Philippines (Opiliones CyphophthalmiStylocellidae) Bulletin of the British Arachnological Society 9 174ndash188

Shear W A and Gruber J (1996) Cyphophthalmid opilionids new toMadagascar two new genera (Opiliones Cyphophthalmi Pettalidae)Bulletin of the British Arachnological Society 10 181ndash186

StaregaW (2008)The second species ofPurcelliaHansenet Soslashrensen 1904(Arachnida Opiliones Pettalidae) from South Africa Polish Journalof Entomology 77 51ndash56

Thorell T (1876) Sopra alcuni Opilioni (Phalangidea) drsquoEuropa e dellrsquoAsiaoccidentale con un quadro dei generi europei de questrsquoOrdine AnnaliMuseo civico Storia naturale Giacomo Doria 8 452ndash508

Manuscript received 23 May 2010 accepted 23 September 2010

Systematic revision of South African pettalids Invertebrate Systematics 401

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 32: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Appendix 1 List of material examined by light microscopy to generate the continuous data set

Parapurcellia peregrinator (Lawrence 1963) ndash holotype lt (NMSA ZA8) from Mariepskop Forest Pilgrimrsquos Rest Mpumalanga South AfricaParapurcellia silvicola (Lawrence 1939) ndash syntype lt (NMSA) from Nkandhla Forest Reserve and Ngoye Forest Kwazulu-Natal South AfricaParapurcellia monticola (Lawrence 1939) ndash syntype lt (NMSA 1838) from Drakensberg Mountains Kwazulu-Natal Province South AfricaParapurcellia rumpiana (Lawrence 1933) ndash syntype lt (NMSA) from Pietermaritzburg Kwazulu-Natal Province South AfricaParapurcellia fissa (Lawrence 1939) ndash syntype lt (NMSA 1119) from Port St Johns Eastern Cape Province South AfricaPurcellia transvaalica Lawrence 1963 ndash syntype lt (NMSA) from Hanglip Forest (near Louis Trichardt) North Transvaal Province South AfricaPurcellia illustrans Hansen amp Soslashrensen 1904 ndash ltlt (ZMUC MCZ) from Table Mountain Western Cape Province South AfricaSpeleosiro argasiformis Lawrence 1931 ndash holotype lt (SAM ENW B001473) from Wynberg Caves Table Mountain Western Cape Province South AfricaRakaia isolata Forster 1952 ndash holotype lt (CM A 14) from North Canterbury South Island New ZealandRakaia lindsayi Forster 1952 ndash holotype lt (MNZ DM 2160) from Stewart Island South AfricaRakaia magna australis Forster 1952 ndash holotype lt (CM A 12) from Lewis Pass South Island New ZealandRakaia media Forster 1948 ndash holotype lt (MNZ DM 2101) from Bay of Plenty North Island New ZealandRakaia solitaria Forster 1948 ndash holotype lt (MNZ DM 2107) from Opouawa Gully West Wairarapa North Island New ZealandRakaia sorenseni digitata Forster 1952 ndash holotype lt (CM A 19) from Chaslands Otago South Island New ZealandAoraki crypta (Forster 1948) ndash holotype lt (MNZ DM 2385) from Coromandel and Bay of Plenty North Island New ZealandAoraki denticula (Forster 1948) ndash holotype lt (MNZ DM 253) from Nelson Marlborough and Buller South Island New ZealandAoraki inerma (Forster 1948) ndash holotype lt (MNZ DM 245) from Te Aroha Mountain Waikato North Island New ZealandAoraki longitarsa (Forster 1952) ndash holotype lt (MNZ DM 2796) from Mount Cook Canterbury South Island New ZealandAoraki tumidata (Forster 1948) ndash holotype lt (MNZ DM 250) from Cuvier Island Coromandel North Island New ZealandKarripurcellia harveyi Giribet 2003 ndash holotype lt (WAM T42519) from Crowea Western Australia AustraliaKarripurcellia peckorum Giribet 2003 ndash holotype lt (WAM T47011) from Warren National Park Western Australia AustraliaKarripurcellia sierwaldae Giribet 2003 ndash holotype lt (WAM T47013) from Warren National Park Western Australia AustraliaAustropurcellia scoparia Juberthie 1988 ndash holotype lt (MNHN) from north-eastern Queensland AustraliaAustropurcellia woodwardi (Forster 1955) ndash holotype lt (QM) from Great Dividing Range Queensland AustraliaChileogovea jocasta (Shear 1993a) ndash holotype lt (AMNH) from Pata de Gallina Region de Biacuteo-biacuteo ChileChileogovea oedipus (Roewer 1961) ndash holotype lt (ZMH) from Isla de Chiloeacute Regioacuten de Los Lagos ChilePettalus lampetides Sharma amp Giribet 2006 ndash holotype lt (MHNG) from Diyaluma Falls Province of Uva Sri LankaPettalus cimiciformis (Pickard-Cambridge 1875) ndash holotype lt (BMNH) from unspecified locality Sri LankaTroglosiro aelleni Juberthie 1979 ndash holotype lt (MHNG) from drsquoAdio Cave Mount Adio New CaledoniaTroglosiro longifossa Sharma amp Giribet 2005 ndash holotype lt (MNHN) from Gicircte Kanua Port Boiseacute New CaledoniaTroglosiro ninqua Shear 1993b ndash holotype lt (MNHN) from Mount Ninqua New CaledoniaHuitaca ventralis Shear 1979 ndash holotype lt (MCZ 14835) from 30Km South of Chinaacutecota Provincia Norte de Santander ColombiaNeogovea sp ndash undescribed lt (MZUSP) from Turuma Mirim State of Amazonas BrazilMetagovea philipi Goodnight amp Goodnight 1980 ndash holotype lt (AMNH) from Los Taxos Cave Morona Santiago Province EquadorCyphophthalmus duricorius Joseph 1868 ndash syntype lt (ZMB 4189) from cave Predjama SloveniaParasiro coiffaiti Juberthie 1956 ndash syntype lt (MNHN) from Girona SpainSiro acaroides (Ewing 1923) ndash syntype lt (USNM) from Coast Range Mountains Benton County Oregon (USA)Siro rubens Latreille 1804 ndash syntype lt (MNHN) from Massif Central FranceSuzukielus sauteri (Roewer 1916) ndash lectotype lt (SMF RI1280) from Yamanaka Suruga JapanStylocellus ramblae Giribet 2002 ndash holotype lt (FMNH) from Botanical Gardens of Singapore SingaporeStylocellus tambusisi Shear 1993c ndash holotype lt (BMNH [E] 1999174) from Tambusisi Mountains Sulawesi Tengah Sulawesi IndonesiaFangensis cavernarum Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Tham Nam Phrathat Si Sawat District Kanchanaburi Province ThailandFangensis spelaeus Schwendinger amp Giribet 2005 ndash holotype lt (MHNG) from Kaeng Lawa Cave in western Thailand

402 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

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Appendix 2 Discrete morphological characters used in the phylogenetic analyses and discussionCharacters new to this analysiswill be discussed in some detail characters used in previous studies are referenced as followsGB indicates character number fromthe matrix analysed in Giribet and Boyer (2002) G indicates Giribet (2003) DG indicates de Bivort and Giribet (2004) BG indicates Boyer and Giribet (2007)

1 Eyes ndash (0) absent (1) present with lens (2) present incorporated into ozophore (GB 2 DG 1 BG 1)2 Ozophore position ndash (0) type 1 (1) type 2 (2) type 3 (3) dorsal facing 45 (GB 2 G 1 DG 2 BG 2)3 Ozophore opening type ndash (0) subterminal (1) infolded (2) terminal with circular opening (3) labial (4) disc-shaped (DG 3 BG 3)4 Ozophore ornamentation ndash (0) absent (1) ornamented on approximately half of its surface (2) entirely ornamented (3) ornamented on less than half of its

surface The surface of the ozophores of most species is entirely covered in tubercular micro-granulate ornamentation (sensu Murphree 1988) of varyingdensityHowever in somespecies in this study (ChileogoveaoedipusandPettalus cimiciformis) roughlyhalf of the surfaceof theozophore lacksornamentation

5 Spiral ornamentation of the ozophore ndash (0) absent (1) present (DG 4 BG 4)6 Proximal end of coxae I meeting along the midline ndash (0) absent (1) present (DG 24 BG 23)7 Second coxae ndash (0) free (1) fused to coxae of legs III (GB 11 G 8 DG 25 BG 24)8 Proximal end of male coxae III meeting along the midline ndash (0) absent (1) present (DG 26 BG 25)9 Coxae II and III endites with processes running along their suture ndash (0) absent (1) present (DG 27 BG 26)10 Gonostome of male in anterior position ndash (0) absent (1) present (DG 29 BG 27)11 Shape of the gonostome ndash (0) semicircular-trapezoidal (1) subhexagonal (2) semicircular (3) oval (4) circular Previous uses of this character (deBivort and

Giribet 2004 Boyer and Giribet 2007) have used just two states semicircular-trapezoidal and subhexagonal Here we recognise that there is substantialdiversity in the shape of the gonostome adding the latter three states Most cyphopthalmids possess semicircular-trapezoidal gonostomes although with thegreater specificity afforded by this extended character state list some species (eg Austropurcellia woodwardiMetagovea philipi and Stylocellus tambusisi)have been recoded as having semicircular gonostomes The only species in this analysis with a different coding under the original scheme was Huitacaventralis with its subhexagonal gonostomeAmong the SouthAfrican pettalids all have either oval or semicircular gonostomes exceptPurcellia illustrans thegonostome of which is semicircular-trapezoidal Generally oval gonostomes are found on the smaller South African pettalids particularly those in the genusParapurcellia

12Anterior projectionsofmale coxae IVenditendash (0) noprojections (1) projections ingonostomewall (2) projectionsadjacent to coxal pore (3) projections alongsuture of coxae IV (BG 29)

13 Endites of coxae IV running adjacent to midline suture for a length longer than gonostome ndash (0) absent (1) present (DG 32 BG 30)14 Oral lappet ndash (0) absent (1) present An oral lappet in the posterior end of the stomotheca is present in Troglosironidae and Neogoveidae but absent or

unevaluated in all other taxa in this study15 Sternum ndash (0) absent (1) present (G 17 DG 28)16 Protruding chelicerae ndash (0) absent (1) present (G 2l BG 5)17 Widest part of cheliceral distal article ndash (0) near base (1) near articulation with mobile digit (DG 5 BG 6) See next character for discussion18 Attenuate chelicerae ndash (0) absent (1) present (GB 3 G 3) Animals often troglomorphic with dramatically tapering second cheliceral segments egHuitaca

ventraliswere originally coded (Giribet andBoyer 2002Giribet 2003) as having lsquoattenuatersquo chelicerae Later (deBivort andGiribet 2004Boyer andGiribet2007) this character state was redefined as the second segment of the chelicerae having its widest part near the base of the article or near the articulationwith themobile digit (character 17 above)All animalswith attenuate cheliceraewere then coded as having thewidest part of the second segment of their chelicerae nearthat articlersquos base Here we recognise that the second segment of the chelicerae of many species is widest near its base but does not have the elongate gracilequality originally intended by lsquoattenuatersquo (eg Chileogovea jocasta) Therefore these two characters have been coded separately Within the South AfricanPettalidae no species have attenuate chelicerae and all but Parapurcellia silvicolaParapurcellia monticola andPurcellia illustrans have chelicerae with thewidest part of the second article near the articulation with the mobile digit

19 Distal segment of chelicerae ornamented ndash (0) absent (1) present (GB 4 DG 6 BG 7)20Dentitionof themobile digit of the cheliceraendash (0) uniform (1) dentition non-uniformwith largest tooth occurring in the centre of the rowof teeth (2) jaggedor

bilobed with smaller lobe distal (GB 6 DG 7 BG 8)21 Basal article of chelicerae with dorsal crest ndash (0) absent (1) present (GB 7 G 5 DG 8 BG 9)22 Basal article of chelicerae with ventral process ndash (0) absent (1) present (GB 8 G 6 DG 9 BG 10)23 Apodeme on chelicerae ndash (0) absent (1) present (BG 11)24Basal article of cheliceraewith a second ventral process ndash (0) absent (1) present In nearly all species thefirst article of the chelicerae has no or a single ventral

process Pettalus lampetides and Stylocellus ramblae have a second ventral process on the basal article of their chelicerae25 Ornamentation of the basal cheliceral segment ndash (0) sparsely ornamented (1) densely ornamented throughout most of its length (2) densely ornamented

centrally The first segment of the chelicerae are densely ornamented in most cyphophthalmids except for in this dataset TroglosiroMetagovea philipi andSiro acaroides which are sparsely ornamented and Siro rubens and Suzukielus sauteri which are densely ornamented centrally

26 Palp trochanter with ventral process ndash (0) absent (1) present (GB 10 G 7 DG 11 BG 12)27Ornamentationof secondpalp articlendash (0) absent (1) less thanhalf ornamented (2)more thenhalf ornamentedThedegreeofornamentationon the secondpalp

article shows considerable variation across and within the cyphophthalmid families In the South African pettalids the second palp article is less than halfornamented except in the small species Parapurcellia amatola and P minuta which have no ornamentation on the third palp article

28 Ornamentation of third palp article ndash (0) absent (1) present Most cyphophthalmids have no ornamentation on the third palp article Within PettalidaeParapurcellia peregrinator Rakaia lindsayi Aoraki tumidata Karripurcellia harveyi K peckorum and Chileogovea oedipus have ornamented third palparticles

29 Solea in tarsus I ndash (0) absent (1) present (GB 12 G 9 DG 13 BG 13)30 Clawof leg Iwithmodifications ndash (0) absent (1) present In a small number of cyphophthalmids the clawof leg I ismodifiedwith lateral pegs or ridges In this

dataset only Suzukielus sauteri hasmodifications to the claw of leg I therefore this character is uninformative with respect to our phylogenetic analyses but isretained in the matrix for reference

31 Claw of leg II with modifications ndash (0) absent (1) present (DG 16 BG 15)

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 403

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 34: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Appendix 2 (continued )

32 Claw of leg II with special row of teeth forming a comb ndash (0) absent (1) present (DG 17 BG 16)33 Claw of leg III with modifications ndash (0) absent (1) present (DG 18 BG 17)34 Claw of leg IV with modifications ndash (0) absent (1) present (DG 19 BG 18)35 Leg II ornamentation ndash (0) all segments smooth (1) metatarsus and tarsus smooth (2) metatarsus partially ornamented and tarsus smooth (3) metatarsus

ornamented and tarsus smooth (4) metatarsus and dorso-basal part of the tarsus ornamented (5) metatarsus ornamented and tarsus almost entirely ornamented(BG 14)

36 Ornamentation of tarsi III and IV ndash (0) absent (1) present (DG 23 BG 22)37 Male tarsus IV ndash (0) entire (1) bisegmented (2) bisegmented dorsally only (GB 15 G 11 DG 21 BG 19)38 Adenostyle ndash (0) lamelliform (1) ending in a tuft of setae (2) fimbriate (3) triangular and heavily sclerotized (4) plumose (5) digitiform (6) bilobed tip

(GB 16 G 12 DG 21 BG 20)39 Adenostyle in the most-basal region of the tarsus ndash (0) absent (1) present (DG 22 BG 21)40Ramblarsquos organndash (0) absent (1) present Inmales ofFangensis (and someother stylocellids) the cuticle of tarsus IV ismodified retrolaterally to formsome sort

of sensorial structure (Schwendinger and Giribet 2005 Clouse et al 2009)41 Spiracle shape ndash (0) circular (1) open circle (2) lsquoCrsquo shaped (GB 21 G 14 DG 33 BG 31)42 Exocrine glands on opisthosoma ndash (0) absent (1) present (BG 32)43 Position of opisthosomal exocrine glands ndash (0) on anterior sternal region (1) on tergite VIII (2) on tergite IX (3) on tergites VIII and IX (4) on tergite IX and

anal plate (BG 33)44Orientation of anal glandopeningndash (0) posterior (1) ventral In cyphophthalmidswith anal glands the opening canbeoriented either ventrally so that it is fully

visible in the ventral view of the animal or posteriorly so that it is visible in the posterior view and only obliquely or tangentially in the ventral view Thischaracter is best assessed by SEMWithin the Pettalidae Parapurcellia Speleosiro Austropurcellia have posterior-oriented anal gland openings PurcelliaChileogovea and Aoraki tumidata have ventral-oriented anal gland openings

45 V-shape modification of sternites 6ndash8 ndash (0) absent (1) present (G 15 DG 36 BG 34)46 Sternite 8 9 and tergite IXndash (0) all free (1) sternites 8 and9medially fused (2) sternite 9 and tergite IX fused but sternite 8 free (3) all fused into corona analis

(4) sternites 8 and 9 completely fused tergite IX free (GB 24 G 16 DG 37 BG 35)47 Relative position of sternite 9 and tergite IX ndash (0) stylocellid type (1) pettalid type (GB 25 DG 38 BG 26)48 Male tergite IX ndash (0) entire (1) bilobed (GB 26 G 17)49 Longitudinal carina in male anal plate ndash (0) absent (1) present (G 18 DG 39 BG 38)50 Ornamentation in midline of male anal plate ndash (0) present (1) absent (DG 40 BG 39)51 Scopulae on anal plate ndash (0) absent (1) present (GB 27 G 20 BG 40)52Positionoforiginof anal scopulaendash (0)posteriormargin (1) anteriormargin (2) posterior ventral surface (3) central ventral surface (4) anteriorventral surface

(BG 41)53 Scopulae on male tergite IX ndash (0) absent (1) present (BG 42)54 Male tergite VIII bilobed ndash (0) absent (1) present (GB 28 G 21 BG 44)55 Large depression onmidline ofmale anal plate ndash (0) absent (depression absent or small) (1) present In animalswith a depressed central region of themale anal

plate this region is either large taking up approximately half ormore of the area of the anal plate (Purcellia andSpeleosiro) or small taking up roughly a third ofthe area of the anal plate (Parapurcellia Rakaia Aoraki longitarsa A tumidata and Austropurcellia woodwardi)

56 Two scopulae originating from each inner margin of tergite VIII ndash (0) absent (1) present (G 22 BG 45)57 Bilobed male anal plate ndash (0) absent (1) present In most pettalids the male anal plate is depressed in the middle or raised on the lateral posterior margins

appearing bilobed This characteristic is absent and the anal plate is entire in Pettalus Chileogovea Karripurcellia Aoraki crypta A denticula A inermaAustropurcellia scoparia and in the members of all other families

58 Anterior opisthosomal sternites of male concave ndash (0) absent (1) present Most cyphophthalmids do not have concave anterior opisthosomal sternitesHowever this character is present in Troglosiro longifossa and T ninqua

59 Male tergite VII bilobed ndash (0) absent (1) present In animals with deeply divided dorsal posterior regions tergite VII can appear bilobed in the dorsal viewThis is absent in most cyphophthalmids but present in Parapurcellia convexa P silvicola Rakaia isolata R lindsayi Aoraki denticula A longitarsa andA tumidata

60 Extension of the scopulae ndash (0) absent (1) present In Aoraki longitarsa and Austropurcellia the scopulae of the anal plate region are greatly extendedappearing as a tail

61 Movable fingers of spermatopositor ndash (0) absent (1) present (DG 45)62 Ovipositor with sense organs ndash (0) absent (1) present (GB 32 DG 46)

404 Invertebrate Systematics B L de Bivort and G Giribet

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 35: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Appendix 3 Continuous morphological characters used in the phylogenetic analysesContinuous characters used in our analysis are either lsquosimplersquo characters (a singlemorphometric ratio or direct measurement) or lsquocomplexrsquo characters comprisingthefirst principal component of twoormore simplecharacters across all taxaSimple characterswere collapsed into a single complex characterwhen independenceanalysis indicated that they were correlated without any outlying examples across all taxa (see methods section) For diagrams of the measurements taken see de

Bivort et al (2010)

1 lsquoOverall sizersquo ndash complex character encoding 49 different raw measurements see de Bivort et al 2010 for full list2 Width between posterior points if the posterior of the animal is bilobed otherwise 03 lsquoProminence of tergite VIIIrsquo ndash complex character encoding the length and concavity of tergite VIII4 Anterior-to-posterior distance from the most posterior to the most anterior points of the posterior margin of tergite VII (tergite VII lsquoconcavityrsquo)5 Concavity of tergite VI6 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites in lateral view7 Distance that the anal plate protrudes distally from the sternites surrounding it in lateral view8 Concavity of sternite 69 Anterior scutum margin concavity10 Distance from the anterior margin of the scutum to the highest dorsal point (dorsal crest) on the first segment of the chelicer11 Length-to-maximum width ratio of the prosoma+ opisthosoma12 lsquoBottom-heavinessrsquo ndash complex character encoding the fractional distance from the anterior to the posterior at which themaximal prosomalwidth andmaximal

opisthosomal width are found13 Fractional distance from the anterior to the posterior of the maximal body thickness14 Fractional distance from the anterior to the posterior of the maximal tergite thickness15 Fractional distance from the anterior to the posterior of the maximal sternite thickness16 Maximal sternite thickness divided by the maximal body thickness17 Maximal tergite thickness divided by the maximal body thickness18 Ratio of the maximal prosomal width to the maximal opisthosomal width19 Minimal width of first tergite divided by the maximal opisthosomal width20 lsquoLaterality of projections of the scutumrsquo ndash complex character encoding the distance from the midline to (one of) the most posterior point(s) divided by the

maximal opisthosomal width the height of the tip of the ozophore divided by the local thickness of the tergite the distance from the prosomal margin to theozophore tips divided by the prosomal width and the distance from the prosomal margin to the ozophore lateral margins divided by the prosomal width

21 Distance from the anterior prosomal margin to the ozophores along the anterior-posterior axis22 Tergite VIII width divided by the width of the opisthosoma23 Maximal length of tergite VIII divided by the length of the scutum24 Maximal length of tergite VII divided by the length of the scutum25 Maximal length of tergite VI divided by the length of the scutum26 Concavity of tergite VIII divided by the length of the scutum27 Concavity of tergite VII divided by the length of the scutum28 Concavity of tergite VI divided by the length of the scutum29 Distance that the anal plate rises dorsally above the lateral-ventral margin of the posterior tergites divided by the maximal body thickness30 Width of the anal plate divided by the maximal opisthosomal width31 Length of the anal plate divided by the total scutum length32 Thickness of the anal plate divided by the total thickness of the scutum33 Length-to-width ratio of the anal plate34 lsquoSize of the anal plate depressionrsquo ndash complex character encoding the width of the central depression of the anal plate divided by the width of the anal plate the

length of the depression divided by the length of the anal plate the width of the unornamented central region of the anal plate divided by the width of the analplate and the fractional area of the anal plate that is modified compared to surrounding (typical) cuticle 0 if these features are absent

35 Width of the anal plate carina divided by the width of the anal plate 0 if carina is absent36 Length of the anal plate carina divided by the length of the anal plate 0 if carina is absent37 Length of the unornamented central region of the anal plate divided by the length of the anal plate 0 if the anal plate is completely ornamented38 Length of the setae on the posterior margin of the anal plate divided by the length of the prosoma39 Length of the setae on the posterior margin of tergite IX divided by the length of the prosoma40 lsquoAnal plate anterior displacementrsquo ndash complex character encoding the maximal length of tergite IX divided by the length of the scutum and the concavities

of sternites 9 8 and 7 each divided by the total length of the scutum41 Maximal length of sternite 9 divided by the length of the scutum42 Maximal length of sternite 8 divided by the length of the scutum43 Maximal length of sternite 7 divided by the length of the scutum44 Maximal length of sternite 6 divided by the length of the scutum45 Width of the fused suture between sternites 8 and 9 divided by the maximal width of the scutum 0 if the sternites are free46 Concavity of sternite 6 divided by the length of the scutum47 Width of the spiracle divided by the maximal width of the scutum48 Distance between the spiracles divided by the maximal width of the scutum49 Degrees of the open portion of the spiracle 0 if the spiracle is circular50 Width of the gonostome divided by the maximal width of the scutum51 Length of the gonostome divided by the length of the scutum

(continued next page )

Systematic revision of South African pettalids Invertebrate Systematics 405

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis

Page 36: A systematic revision of the South African …debivort.org/pdf/Revision_of_the_South_African...A systematic revision of the South African Pettalidae (Arachnida:Opiliones:Cyphophthalmi)

Appendix 3 (continued )

52 Length-to-width ratio of the gonostome53 lsquoSize of the sternal regionrsquo ndash complex character encoding the distance from the anterior margin of the scutum to the centre of the gonostome divided by the

length of the scutum the distance from the anteriormargin of the gonostome to the posteriormargin of the palpal enditesmouthparts divided by the length of thescutum and the length of the suture between the left and right coxae IV endites at the midline divided by the length of the scutum

54 Distance along the anterior-posterior axis from the centre of the gonostome to the lateral margin of sternite 1 divided by the length of the scutum55 Length of the suture between the left and right coxae I endites at the midline divided by the length of the scutum56 Length of the suture between the left and right coxae II endites at the midline divided by the length of the scutum57 Length of the suture between the left and right coxae III endites at the midline divided by the length of the scutum58 Length of the suture between the coxae II and III endites divided by the width of the prosoma59 Maximal width across the coxal endites divided by the width of the prosoma60 lsquoLength of anterior coxaersquo ndash complex character encoding the length of coxa 1 measured from the its most medial point of its endite to its most distal point

(lsquoradial lengthrsquo of coxae I) divided by the width of the prosoma and the radial length of coxae II divided by the width of the prosoma61 lsquoLength of posterior coxaersquo ndash complex character encoding the radial length of coxae III divided by the width of the prosoma and the radial length of coxae IV

divided by the width of the prosoma62 Distance between the tips of coxa I and II divided by the length of the scutum63 Distance between the tips of coxa II and III divided by the length of the scutum64 Distance between the tips of coxa III and IV divided by the length of the scutum65 Thickness of coxae IV divided by the maximal thickness of the body66 Distance between the lateral margins of coxae IV divided by the width of the prosoma67 Distance along the anterior-posterior axis from the most anterior point of the prosoma to the margin of the prosoma at the midline divided by the length of the

scutum68 Length of the first segment of the chelicer divided by the length of the scutum69 Length of the second segment of the chelicer divided by the length of the scutum70 lsquoLength of themobile digit of the chelicerrsquo ndash complex character encoding the length of themobile digit of the chelicer divided by the length of the scutum and

the length of the mobile digit of the chelicer divided by the length of the second segment of the chelicer71 Maximal width of the first segment of the chelicer divided by the width of the prosoma72 Maximal width of the second segment of the chelicer divided by the width of the prosoma73 lsquoFirst segment of the chelicer thicknessrsquo ndash complex character encoding the maximal thickness of the first segment of the chelicer divided by the maximal

thickness of the body and the minimum thickness of the first segment of the chelicer just proximal to the dorsal crest divided by the maximal thickness of thebody

74 Height of dorsal crest above the point proximal to the dorsal crest where the first segment of the chelicer attains its minimal thickness divided the maximalthickness of the first segment of the chelicer

75Distance from the proximal endof thefirst segment of the chelicer to thehighest point of thedorsal crest dividedby the length of thefirst segment of the chelicer76 P to D position of maximal width of cheliceral segment 277 Distance along the midline from the anterior margin of the scutum to the dorsal crests divided by the length of the scutum78 Length overwhich the second segment of the chelicer is ornamented divided by the length of the second segment of the chelicer 0 if the second segment of the

chelicer is unornamented

406 Invertebrate Systematics B L de Bivort and G Giribet

httpwwwpublishcsiroaujournalsis


Systematic Innovation

Systematic Searching for Systematic Reviews

Updated - What makes systematic review systematic - Anna Sidorchuk

A new genus of cyphophthalmid from the Iberian Peninsula ...debivort.org/pdf/...from_the_Iberian_Peninsula.pdf · The cyphophthalmid fauna of the Iberian Peninsula is not extraordinary

Systematic Innovation of Products, Processes, and Servicescdn.executive.mit.edu/.../systematic-innovation-of-products-processe… · Systematic Innovation of Products, Processes,

Epinephrine in cardiac arrest: systematic review and · PDF fileEpinephrine in cardiac arrest: systematic review and meta-analysis. ... systematic review and meta-analysis ... systematic

SYSTEMATIC REVIEW A systematic review and meta-analysis

MSN Systematic Evaluation Plan Systematic Evaluation Plan ...MSN Systematic Evaluation Plan Systematic Evaluation Plan for Standard One I. The mission of the nursing education unit

SIMMONS- SYSTEMATIC THEOLOGY CONTENTS Books II/Simmons - A Systematic Study of... · SIMMONS- SYSTEMATIC THEOLOGY CONTENTS A Systematic Study of Bible Doctrine BY Thomas Paul Simmons

Mapping the systematic literature studies about so …...On the other hand, the terms related to systematic literature reviews are: SLR, Systematic Literature Review, systematic mapping,

A molecular phylogeny of the temperate Gondwanan family ...  · Web viewA molecular phylogeny of the temperate Gondwanan family Pettalidae (Arachnida, Opiliones, Cyphophthalmi) and

Systematic Romantic

A molecular phylogeny of the temperate Gondwanan …boyer/Site/Publications_files/Giribet...family Pettalidae (Arachnida, Opiliones, Cyphophthalmi) and the limits of taxonomic

Systematic Min

Systematic Reviews. What is systematic review? - Well documented of intervention research -Scientific methodology -reduced systematic errors (biases)

Systematic Searching Systematic Reviews · Systematic Searching Systematic Reviews Literature search Tomas Allen ... and other systematic reviews, clinical trials, and more. Cochrane

Systematic Mapping Studies - Uni Koblenz-LandauMarcel Heinz Systematic Mapping Studies 23. Juli 2014 3 / 44 Systematic Review : History Creating systematic reviews is an established

Systematic Literature Reviews - GitHub Pages · Systematic literature review Systematic mapping review Systematic scoping review Situates a study within the relevant literature, non-systematic

Systematic risk

Social context modulates idiosyncrasy of behaviour in the ...debivort.org/pdf/Social_context_modulates_idiosyncrasy.pdf · cockroach Blaberus discoidalis James D. Crall a, b, *,1,

A morphometrics-based phylogeny of the temperate …debivort.org/pdf/A_morphometrics-based_phylogeny_of_the_Pettalidae.pdfRowland Institute at Harvard, Department of Organismic and

Systematic Transfer Plan/ Systematic Withdrawal Plan (Form 3)€¦ ·  · 2017-12-074.2 Systematic Transfer Plan ... Systematic Transfer Plan/ Systematic Withdrawal Plan (Form 3)

SYSTEMATIC REVIEW Open Access Guidelines for guideline developers: a systematic … · 2017-04-06 · SYSTEMATIC REVIEW Open Access Guidelines for guideline developers: a systematic

“THE SYSTEMATIC STUDY OF HUMAN SOCIETY” Systematic

Systematic review

Systematic Theology - The Cambron Institutethecambroninstitute.org/library/Systematic_Theology.pdf · systematic theology c:\documents and settings\jboehm\desktop\systematic theology.doc

A Systematic Review of Systematic Review Process Research ... Kitchenham - A systematic revie… · Keywords: systematic review; systematic literature review; systematic review methodology;

Systematic Desensitization

A molecular phylogeny of the temperate Gondwanan family ...debivort.org/pdf/A_molecular_phylogeny_of_Pettalidae.pdf · A molecular phylogeny of the temperate Gondwanan family Pettalidae

SYSTEMATIC REVIEW Open Access Toward systematic reviews …

Systematic approach

Systematic Sampling

Systematic reviews: making them policy relevant · A systematic review is a review of research literature using methods that are systematic, explicit and accountable2. Systematic

Systematic Literature Review - Poltekkes Pontianak · • Campbell collaborationfor systematic reviews of social interventions • Cochrane collaborationfor systematic reviews of

INVESTMENT CLASSIFICATION PROJECT: Systematic Valued1pvbs8relied5.cloudfront.net/.../Systematic-Value... · INVESTMENT CLASSIFICATION PROJECT: Systematic Value MARKET INSIGHT PAPER