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© The Norwegian Academy of Science and Letters 2006 • Zoologica Scripta,

35

, 1, January 2006, pp19–62

19

Kuntner, M. (2006) Phylogenetic systematics of the Gondwanan nephilid spider lineageClitaetrinae (Araneae, Nephilidae). —

Zoologica Scripta

,

35

, 19–62.This study revises the taxonomy, biology, phylogeny, and biogeography of the basal-mostnephilid spider lineage, the Clitaetrinae, with the least known nephilid genus

Clitaetra

. Thefive previously known species are redescribed:

Clitaetra clathrata

Simon from western Africa,

C. simoni

Benoit from central Africa,

C. episinoides

Simon from the Comoro Islands and Mayotte,

C. perroti

Simon from Madagascar, and

C. thisbe

Simon from Sri Lanka with first descriptionsof the males of

C. clathrata

and

C. perroti

. Additionally,

C. irenae

sp. nov. is described in bothsexes from southern Africa.

Clitaetra

biology, so far largely unknown, is presented here basedon observations of

C. irenae

in South Africa, and clitaetrine anatomy is summarized to assessphylogenetic homologies. A species-level phylogenetic analysis of 32 taxa scored for 197morphological and behavioural characters results in eight most parsimonious cladograms andplaces

Clitaetra

as sister to the clade (

Herennia

+ (

Nephilengys

+

Nephila

)). Thus, the orb-weavingspider family Nephilidae Simon contains the (sub)tropical genera

Nephila

,

Nephilengys

,

Herennia,

and

Clitaetra

, but not

Deliochus

or

Phonognatha

.

Contra

recent cladistic treatments, the nephilinesare not tetragnathids, but the sister group to the newly proposed clade, Nephilidae, is ambigu-ous. The three species clades (subgenera) within

Clitaetra

show a seemingly old Gondwananbiogeographic pattern:

Afroetra

subgen. nov., with the three mainland African species, is sisterto

Clitaetra

with the two Indian Ocean island species.

Indoetra

subgen. nov. contains theunstudied species from Sri Lanka,

C. thisbe

. Future understanding of the morphology and bio-logy of

C. thisbe

is important for the polarization of many nephilid features. Vicariancewould estimate the clitaetrine subgeneric clades and basal nephilid lineages to be at least 160Myr old and of Gondwanan origin.

Matja

°

Kuntner, Department of Entomology, National Museum of Natural History, SmithsonianInstitution, Washington, D.C. 20013–7012, USA and Department of Biological Sciences, GeorgeWashington University, Washington, D.C. 20052, USA. E-mail: [email protected]*Present address: Institute of Biology, Scientific Research Centre of the Slovenian Academy of Sciencesand Arts, Novi trg 2, P.O. Box 306, SI-1001 Ljubljana, Slovenia.

Blackwell Publishing Ltd

Phylogenetic systematics of the Gondwanan nephilid spider lineage Clitaetrinae (Araneae, Nephilidae)

M

ATJA

Z

K

UNTNER

*

Accepted: 13 September 2005doi:10.1111/j.1463-6409.2006.00220.x

Introduction

The orb-weaving spider family Nephilidae Simon containsthe (sub)tropical genera

Nephila

Leach, 1815,

Nephilengys

L. Koch, 1872,

Herennia

Thorell, 1877 and

Clitaetra

Simon,1889 (Kuntner 2005, this study). Perhaps the best knownnephilid features are the large ‘golden’ orb-webs of

Nephila

,reaching 1.5 m in diameter (own data), the extreme sexualsize dimorphism of

Nephila

and

Nephilengys

in which themales are a fraction of the size of the females (Coddington

et al

. 1997; Hormiga

et al

. 1995, 2000; Harvey

et al

. in press;Kuntner 2005), and the bizarre sexual biology of

Nephila

and

Herennia

, where broken-off male sperm transferring organsare commonly found stuck in female genital openings (Kuntner2005). In contrast, little is known about

Clitaetra

diversityand less about its biology.

Nephilinae was first recognized by Simon (1894), but itsfamilial placement within Araneoidea has changed repeatedly(see also Kuntner 2002, 2005; Kuntner & Hormiga 2002).Simon’s (1894) argiopid subfamily Nephilinae consisted ofthe genera

Singotypa

Simon, 1894,

Phonognatha

Simon, 1894;

Deliochus

Simon, 1894,

Nephila

(

Nephilengys

included),

Clita-etra

, and

Herennia

. Later,

Singafrotypa

Benoit, 1962 and

Perilla

Thorell, 1895 were included in nephilines, and

Singotypa

wassynonymized (Dondale 1966) with

Phonognatha.

For most ofthe ‘precladistic’ 20th century Nephilinae remained placedwithin Araneidae. Levi (1980) placed

Nephila

in the araneidsubfamily Metinae. Later, Levi (1986: 99) considered

Nephila

and

Nephilengys

to ‘probably’ belong in Tetragnathidae andLevi & von Eickstedt (1989) retained the whole subfamilyNephilinae in Tetragnathidae.

Nephilid spider lineage Clitaetrinae

•

M. Kuntner

20

Zoologica Scripta,

35

, 1, January 2006, pp19–62 • © The Norwegian Academy of Science and Letters 2006

The first cladistic analysis containing nephiline genera(Coddington 1990), with 32 orbicularian taxa scored for87 morphological and behavioural characters, recovered amonophyletic clade (

Nephila

and

Nephilengys

) as sister totetragnathid genera

Meta

C

.

L. Koch, 1836,

Leucauge

White,1841,

Tetragnatha

Latreille, 1804 and

Glenognatha

Simon, 1887.The nephiline placement of

Phonognatha

and the monophylyof Nephilinae and Tetragnathidae were further tested byHormiga

et al

. (1995) with a denser taxonomic sample. Theresults of the latter study corroborated Coddington’s (1990)hypothesis of Nephilinae as the sister group of the remainingtetragnathid lineages and hypothesized the relationships ofnephiline genera: (

Phonognatha

(

Clitaetra

(

Nephila

(

Herennia

+

Nephilengys

)))). The study of Hormiga

et al

. (1995), based onmorphological and behavioural data, recovered the mono-phyly of Nephilinae and of Tetragnathidae, a result later cor-roborated by Griswold

et al

. (1998) within a broad taxonomicsample of araneoids. Kuntner & Hormiga (2002) and Kunt-ner (2002) revised the ‘nephiline’ genera

Singafrotypa

and

Perilla

, and placed them phylogenetically in Araneinae(Araneidae). Recent studies (Pan

et al

. 2004; Wunderlich 2004)including a preliminary finding from this research (Kuntner2003) have hypothesized nephilines as sister to Araneidaerather than Tetragnathidae. However, no prior work hasemphasized

Deliochus

and no nephiline phylogeny exists atthe species level, making evolutionary interpretations of theavailable biological data difficult or impossible.

Taxonomic history

Simon (1889) erected the monotypic genus

Clitaetra

(thenin Epeiridae) and described

C. episinoides

Simon, 1889 fromMayotte. In his encyclopedic treatise of world spiders, Simon(1894) redescribed the genus (original misspelling

Clytaetra

)with a single drawing of the ocular area of

C. episinoides

(typespecies) and described a new species,

Clitaetra perroti

fromthe eastern Madagascar coast. Simon (1894) placed

Clitaetra

in Clitaetreae within Nephilinae, family Argiopidae. In 1903,Simon described the third species,

Clytaetra

[sic]

thisbe

fromSri Lanka, which remains the only

Clitaetra

species foundoutside Africa with Madagascar. Simon (1907) then described

C. clathrata

Simon, 1907 from Guinea Bissau (Guinee portug.),and diagnosed all four species. Benoit (1962) redescribed

C. clathrata

and described a new species,

Clitaetra simoni

from Zaire (Sankuru, D.R. Congo). Schmidt & Jocqué (1986)described and illustrated the male of

C. episinoides

from theComoro Islands. Hormiga

et al

. (1995: 330) illustrated the malepalps of

C. perroti

(misidentified as

C. episinoides

) (theirfig. 9D,E) and

C. clathrata

(as

Clitaetra

sp.) (their fig. 9A–C)and provided spinneret images of

C. irenae

sp. nov. (as

Clita-etra

sp.; their fig. 15A–D).Simon (1889, 1894, 1903, 1907) provided brief descrip-

tions in Latin, without illustrations, based exclusively on

scarce female material and did not designate types. Un-surprisingly, most species have remained unstudied and theiridentification is difficult. In this study,

Clitaetra

species areredescribed, diagnosed and illustrated using type specimens,with first descriptions of the males of

C. perroti

and

C. clath-rata

. A new species,

C. irenae

sp. nov. from southern Africa isdescribed with extensive information on its biology.

Scope

Clitaetra

specimens are rare in collections and

Clitaetra

naturalhistory has never been described. A photograph of

Clitaetra

in Preston-Mafham (1991 : 117) (=

C. irenae

sp. nov.) suggestedarboricolous habits with a web against tree trunks (confirmedby personal communications of Astri Leroy (on

C. irenae

sp.nov.) and of Rudy Jocqué (on

C. episinoides

)), similar to

Herennia

(Robinson & Lubin 1979; Hormiga

et al

. 1995;Kuntner 2005). Limited behaviour entries coded for

Clitaetra

in a phylogenetic analysis of tetragnathid genera byHormiga

et al

. (1995) demonstrate the need for new materialand behavioural data. I focused on the behaviour of thenew species from South Africa (

C. irenae

sp. nov.) during anexpedition to KwaZulu-Natal in 2001.

The present paper, the first of the nephilid taxonomic treat-ments (see also Kuntner 2005) provides detailed methodsused in nephilid taxonomy and behavioural research, andrevises the biology and the taxonomy of

Clitaetra

.

Clitaetra

morphology and behaviour are summarized for use in nephilidphylogeny and for a phylogenetic definition of the genus.A species-level phylogeny is devised for

Clitaetra

and selectednephilid and araneoid outgroups.

Clitaetra

is sister to all othernephilid genera, and thus its biology, when better known,may alter the homology and polarization of many nephilidfeatures as understood today. The phylogenetic resultssuggest that

contra

to recent cladistic studies (Coddington 1990;Hormiga

et al

. 1995; Scharff & Coddington 1997; Griswoldet al. 1998) nephilines are not tetragnathids, and thus areproposed to be treated as a family.

Materials and methodsNephilid specimen databaseThe nephilid specimen database (available for all nephilidgenera; see Kuntner 2005) was developed in BIOTA 1.6.0(Colwell 1999). At least one specimen from each examinedsample received a unique specimen code (a required field inBIOTA), which consisted of the two letter genus code (‘cl’for Clitaetra, ‘ne’ for Nephila, ‘ng’ for Nephilengys, ‘he’ forHerennia) followed by the sample number, a letter indicatinggender or stage (‘f’ for a female, ‘m’ for a male, ‘j’ for a juvenile)and the consecutive number within sample. Other specimendata included species identification and date, gender, abundance,type status, museum depository (see Museum abbreviations),and optional notes. The locality data entered in BIOTA for

M. Kuntner • Nephilid spider lineage Clitaetrinae

© The Norwegian Academy of Science and Letters 2006 • Zoologica Scripta, 35, 1, January 2006, pp19–62 21

each sample, if known, were: a unique locality code, localityname, district, state/province, country, elevation, latitudeand longitude and their accuracy (by convention, the word‘approx.’ in this field means the approximate coordinateswere found later through maps or gazetteers, not measuredon spot). If interpreted, the original spellings were includedin square brackets. The collection data entered in BIOTA foreach sample, if known, were: a unique collection code, collectedby (in case of one collector the first name abbreviation followedby full last name, in case of two collectors both last namesseparated by ‘&’, and in case of several collectors the last nameof the first collector followed by ‘& al.’), date collected, site(if available) and source (‘label’ indicating museum label and‘notes’ as personal notes from the field).

The nephilid specimen database, currently containing morethan 4000 entries, is available from the author. Upon publica-tion of all four nephilid revisions (Kuntner in prep., 2005, thisstudy ), the database will be made available on the internet.

Behavioural observationsAll behavioural observations were carried out in the field inthe spiders’ natural environment; voucher specimens werecollected and deposited at USNM. Observations were doneduring the day and at night, when a red filtered headlamp wasused. A microcassette recorder along with a field notebookwas used to document behaviours. Behaviours and web archi-tecture were photographed after the webs were dusted withcornstarch (Eberhard 1976; Carico 1977). Additionally,samples of crucial parts of orb webs were taken using micro-scope slides with double-sided adhesive tape on the edges.These samples were later examined under the microscope toestablish exact thread junctions and their nature.

Some behaviours are described here for the first time, andmany are further discussed in other nephilid phylogenetictreatments (Kuntner 2005; Kuntner, Hormiga and Codding-ton unpublished). Most behaviours, however, have beendescribed and discussed in the literature (for review see Eber-hard 1982) and used in recent phylogenetic analyses usingorbicularian spiders as terminals (Coddington 1986a,b,c,1990; Hormiga et al. 1995; Scharff & Coddington 1997;Griswold et al. 1998; Agnarsson 2003, 2004, 2005). Thenomenclature used here attempts to follow the publishedaccounts (see Abbreviations).

Morphological examinationMost morphological observations and illustrations of exter-nal structures were made using a Leica MZ APO dissectingmicroscope with a camera lucida. For internal palpal andepigynal anatomy a compound microscope Leica DMRMwith a camera lucida was used. Microscope images were takenusing a Nikon DXM 1200 digital camera, and assembledwith Syncroscopy Automontage software. Digital SEM photo-

graphs were taken on a Leo 1430VP scanning electronmicroscope at the Department of Biological Sciences,George Washington University. For SEM preparation, speci-mens were cleaned ultrasonically for 1 min, transferred to100% ethanol overnight, dissected and submitted to criticalpoint drying, mounted on rivets using glue and copper wire,then sputter-coated.

Female genitalia and male palps were excised using scalpelsand needles. Male palpal anatomy (trajectory of the ducts)and internal female genitalic structure were examined byclearing the organs in methyl salicylate (Holm 1979), mount-ing them on a temporary slide (Coddington 1983) andillustrated under compound microscope. In spiders withlarge and heavily sclerotized internal genitalia (Nephila,Nephilengys) this technique works poorly. Thus, in order todecipher the trajectory of the copulatory ducts, the epigynawere exposed to concentrated KOH to completely digest anysoft tissues. Cleared genitalia were illustrated under a dissect-ing microscope and sometimes further dissected to exposethe details. With this technique it is often necessary to inter-pret the relationships of the positions where the copulatoryand fertilization ducts attach to the spermatheca lumen. Malepalps were expanded by exposing them to concentrated KOHfor up to an hour, followed by immersion in distilled water,and the process repeated as needed. Nephilid palps rarelyexpand fully, probably due to relatively small haematodochae,and the extent and the close association between the long embolusand the grooved embolic conductor. Often, the embolusneeded to be pulled out of the embolic conductor, riskingbreakage of structures. To fully understand the sclerite asso-ciations, more destructive dissections have to be made, oftenatomizing the palps into separate sclerites and their parts.

All measurements are in millimeters and were made usinga micrometer eyepiece. Prosoma and opisthosoma length andheight were measured in lateral view, the width in dorsalview; all measured at widest points (prosoma height meas-ured at head region). Eye widths are maximum diameters ofthe eye lens and eye separations were measured betweenlenses. Leg segments were measured in detached legs. Onlyjoints distal from the trochanter were measured, as thedetached leg breaks between the trochanter and the femur.Maximum lengths of femur, patella, tibia, metatarsus andtarsus are reported; the total of these is the approximated totalleg length [if all leg joints were measured at the maximumjoint leg, the sum would overestimate the total leg lengthbecause joints partly overlap at junctions].

ImagingLarge nephilid females commonly exhibit diagnostic somaticmorphology and colour patterns. Less conspicuous, notablyin Nephila and Clitaetra, is the epigynal morphology, oftencovered with the tissue posterior to the epigynal plate. Thus,

Nephilid spider lineage Clitaetrinae • M. Kuntner

22 Zoologica Scripta, 35, 1, January 2006, pp19–62 • © The Norwegian Academy of Science and Letters 2006

female habitus illustrations in addition to the epigynum areprovided for all species, with variation if applicable. In con-trast to the large females, the male somatic morphology is fairlyuniform within genera. Their (unexpanded) palps, however,are diagnostic, and thus illustrated for all species. Ectal viewsare especially useful and should suffice for species identifica-tion (provided the examined palp is viewed from exactly thesame angle, for which it is best detached from the spider).

Pencil illustrations via camera lucida were kept on acid-free archival paper. Simple line illustrations were inked onVellum film; shaded illustrations were inked on fine grainedcoquille board and further rendered with black, soft Prisma-color pencils. All nondigital artwork (photographic slidesincluded) was scanned for digital manipulation in AdobePhotoshop 7.0, where digital microscope and SEM imageswere further manipulated. All image plates were assembledand labelled in Adobe Illustrator 10.

Phylogenetic analysisAll known Clitaetra species represented the ingroup. Nephili-nae sensu Hormiga et al. (1995) included the genera Nephila,Nephilengys, Herennia, Clitaetra and Phonognatha, and weresister to other tetragnathids. I used the type species ofall nephiline genera in the analysis: Phonognatha graeffei(Keyserling, 1865), Herennia multipuncta Doleschall, 1859,Nephilengys malabarensis (Walckenaer, 1842) and Nephilapilipes (Fabricius, 1793). I included the second species ofNephilengys (N. cruentata (Fabricius, 1775)) and Herennia(H. papuana Thorell, 1881) to test for the generic monophyly.In the large pantropical genus Nephila more exemplars wereused. Harvey et al. (in press.) revised the five Australasianspecies, of which four are represented in this analysis: N. pilipes,N. antipodiana (Walckenaer, 1842), N. plumipes (Latreille,1804) and N. edulis (Labillardière, 1799). Additional speciesfrom the Americas (N. clavipes (Linnaeus, 1767)), Africa(N. fenestrata Thorell, 1859), and Asia (N. clavata L. Koch,1878) were included. About six Australasian species ofDeliochus are known (two described). The particular speciesincluded here, for which I have behavioural data, is un-described. The non-nephiline outgroup taxa are a subset ofthose in Kuntner, Hormiga and Coddington (unpublished;see their specimen information): Deinopis MacLeay, 1839 andUloborus Latreille, 1806 are the nonaraneoid outgroups. Ara-neids are represented by Araneus Clerck, 1757 and two ArgiopeAudouin, 1826 species with remarkably different somaticand genitalic morphology. Tetragnathids are represented byTetragnatha, Meta and Leucauge, and the reduced piriformclade taxa are as in Hormiga et al. (1995): Linyphia Latreille,1804 (Linyphiidae), Pimoa Chamberlin & Ivie, 1943 (Pimoi-dae), Steatoda Sundevall, 1833 (Theridiidae), NesticusThorell, 1869 (Nesticidae), and Epeirotypus O. P.-Cambridge,1894 (Theridiosomatidae).

Appendix 1 lists the characters and character states used inthe analysis. The characters and their homologies will be dis-cussed in more detail elsewhere (Kuntner 2005; Kuntner,Hormiga and Coddington unpublished). The phylogeneticmatrix (Appendix 2) of 197 characters scored for 32 taxa inNONA format (ss) is available from the author and will besubmitted to TreeBASE (www.treebase.org).

For the cladistic analysis I used NONA version 2.0 (Golo-boff 1993) under both amb – (default) and amb = (allows formore ambiguous support) and with parameters hold 1000(keeps 1000 trees during searches), mult*500 (random taxonaddition), max* (performs the tree bisection-reconnection(TBR)), and sswap (branch swapping algorithm). I usedWinclada 1.00.08 (Nixon 2002) to display and manipulatetrees and matrices for NONA. Successive character weight-ing (Farris 1969) analysis was performed in NONA with thecommand run swt.run hold10000 hold/100 mult*100(using the macro swt.run). The bootstrap values (Felsenstein1985) were calculated in Winclada using default settings (100replications, mult*10). Bremer support or decay index values(Bremer 1988, 1994) were calculated in NONA using thecommand bs10 and hold 100000.

All trees are output from Winclada and their formatdoes not imply nonmonophyly of the group Deinopoidearepresented here by the two primary outgroups, Deinopis +Uloborus.

ClassificationI use a compromise approach between the (classical) zoo-logical nomenclature (ICZN 1999) and phylogenetic nomen-clature (Cantino & de Queiroz 2004). A review of the debatewith a rationale for such combined classification is in prepa-ration (see also Kuntner 2005). Clade names in this study areconsistent with zoological ranks up to the family level, but areprecisely circumscribed following the PhyloCode articles 7,9–11 (Cantino & de Queiroz 2004). Phylogenetic definitions(PC Article 9, Note 9.4.1) used here are node-based where‘clade (A and B)’ means the least inclusive clade containingA and B. Due to ambiguous character support (Fig. 28),apomorphy-based clade definitions or combined definitionsare not used. Because the PhyloCode has not yet taken effectand the registration database has not yet been implemented,the names proposed here are not registered (Article 8).

Museum abbreviationsCAS: California Academy of Sciences, San Francisco, USA;MCSNG: Museo Civico di Storia Naturale, Genova, Italy;MCZ. Museum of Comparative Zoology, Harvard Univer-sity, Cambridge, USA, MNHN: Muséum national d’histoirenaturelle, Paris, France; PPRI: Plant Protection ResearchInstitute, Pretoria, South Africa; RMCA: Musée Royal del’Afrique Centrale, Tervuren, Belgium; USNM: National



Museum of Natural History (former United States NationalMuseum), Smithsonian Institution, Washington, DC, USA.

Anatomic and behavioural abbreviationsAC: aciniform gland spigot(s); AG: aggregate gland spigot(s);ALS: anterior lateral spinneret; AT: anal tubercle; BH: basalhaematodocha; BL: booklung(s); CB: cymbium; CD: copula-tory duct; ChD: cheliceral denticles; CO: copulatory open-ing; CY: cylindrical gland spigot(s); E: embolus; EAD:epigynal anterior depression; EB: embolus base; EC: embolicconductor; ECG: embolic conductor groove; ELC: epigynallateral curve; EP: epigynum; ES: epigynal septum; ETm:embolus-tegulum membrane; F: fundus; FD: fertilizationduct; FL: flagelliform gland spigot(s); M: membrane (mem-branous); MAP: major ampullate gland spigot(s); mAP:minor ampullate gland spigot(s); N: nubbin; NSS: nonstickyspiral (also termed auxiliary or temporary spiral); P: paracym-bium; PCT: promarginal cheliceral teeth; PI: piriform glandspigot(s); PLS: posterior lateral spinneret; PMS: posteriormedian spinneret; PS: paracymbial seta(e); PPS: palpal patel-lar seta(e); PSL: prosomal supracheliceral lobe; RCT: retro-marginal cheliceral teeth; S: spermatheca; Sc: scutum; SD:sperm duct; SS: sticky spiral; ST: subtegulum; T: tegulum;TO: tarsal organ; Tr: trichobothrium(a); VTA: ventral tegularapophysis.

ResultsBiologyI studied Clitaetra irenae sp. nov. in South Africa at FaniesIsland, Sodwana Bay, Tembe Elephant Park, and PhindaResource Reserve (photographs available on request). Thespecies is abundant in sand forests of northern lowlandKwaZulu-Natal. The spiders typically spin their orb webs ontree trunks, but one female was taken from a web built on arock outcrop at Phinda, and spiders were commonly collectedfrom wooden walls at localities around St. Lucia Estuary. Thewebs (photographs available on request) could be describedas ladder-webs, as they are built more or less vertically, areelongated with parallel vertical sides and their ‘spirals’ runparallel. The webs are typically built within tree trunkdepressions such that the clearance between the web and thetree is greatest at the hub (typically 2–4 cm in female webs).The hub of the orb is reinforced with a fine silk mesh, whichmay be homologous to the ‘hub-cup’ of Herennia and to theretreat of Nephilengys. The spider rests at the hub, typicallywith prey remains suspended above the hub.

The webs of adult females are eccentric, the lower portionbeing larger than the upper (the ratio top frame to hub/topto bottom frames ranged from 0.21 in a female at Phinda to0.31 in female at Makakatana); only some juvenile webs areeccentric. Females, juveniles and even adult males were seenin prey-ready position head-down at the hub, with legs I and

II partially flexed. I collected three males from their ownminiature orb webs (photographs available on request), and asample from Phinda contains no sticky silk, suggesting theweb was made after the last moult. Adult males were com-monly found on webs of either adult or juvenile females anda male was observed guarding a female, which had alreadydeposited her eggs, with the egg-sacs suspended in the planeof the web above the hub.

Clitaetra irenae builds webs at night and in the early morn-ing hours, sometimes only partially renewing the old web(typically, the hub and an orb sector are retained and mayonly be reinforced). Most specimens observed at FaniesIsland (n = 16) did not build between 05:00 and 01:00 exceptwhen fixing partially damaged webs, but one female at Tembewas doing frame behaviour at 20:30. Typically, web-buildingbehaviour started at around 02:00 and was terminated by05:00.

The following summarizes web architecture and buildingbehaviour (for character descriptions see Kuntner 2005):sticky spiral localization is done with outer fourth leg; hubloop-NSS transition is gradual; hub bite-out is absent; hub isalways closed; radii are attached twice on the frame (exceptlate radii, attached singly), and once on the hub; radii are notcut-and-reeled; nonsticky spiral (NSS) persists in finishedweb [NB: This behaviour was observed as it occurred (n = 2)and was confirmed from the web samples of adults and juve-niles. The persisting NSS are few and far between comparedto Nephila, and often get glued to the SS. Hence, observationsfrom the finished webs and from photographs repeatedlyfailed to establish the presence of NSS in finished webs];barrier webs are absent; NSS form is not zig-zag; silk colouris white (unlike in Nephila, where it is golden); secondary andtertiary late radii are numerous, originating on NSS; hub silkenforcement is present.

When disturbed, females and juveniles responded eitherby shaking their body or rushing to the other side of the orb.The former behaviour was common and would last up toa minute. The latter behaviour (photographs available onrequest), involves the spider smoothly and rapidly movingfrom the outer to the inner side of the orb (apparentlythrough the radii), where it rests protected by the web on oneside and the tree on the other, and showing its venter throughthe hub. This behaviour only takes a fraction of a second; asimilar sequence is known to occur in Herennia, some Nephilaand the araneid Argiope (Kuntner 2005). Females guardedtheir egg-sacs suspended in the web between the hub and theupper frame and did not abandon their protective positioneven in a highly damaged web and under stress.

Attack behaviour depended on the size of prey, but neverinvolved attack wrapping. Spiders attack small prey (flies) bybiting, followed by immediate digestion and consumption(n = 5). They attack larger prey (moth, n = 2) with a bite first,



wrap second, followed by the spider hanging the prey underthe hub, and assuming rest posture at the hub. This seems tobe a stereotypical behaviour, as it was immediately followedby re-grabbing the suspended prey and feeding on it. Asimilar behavioural sequence is also known in Nephila andNephilengys (own data).

Predatory mimetid spiders invade the webs of Clitaetra(photograph available on request). If the spiders were forcedto abandon their webs, their predators would include antsthat are abundant on tree trunks. No symbionts wereobserved in C. irenae webs. A female of Rugathodes sp.(Theridiidae) was taken from a female C. irenae web fromTembe (along with a Clitaetra male). The genus is probablynot a kleptoparasite of Clitaetra.

Web-building behaviour and natural history of all otherClitaetra species remains unstudied. A photograph ofC. episinoides from Comoro Islands (by R. Jocqué) suggeststhat the species biology may be similar to that of C. irenae inthat the species makes its orb web with a ‘hub-cup’ on treetrunks and the prey-ready position is at the hub with legs Iand II partially flexed.

PhylogenyEqually weighted analyses resulted in eight most parsimoni-ous trees (L = 531, CI = 42, RI = 70). In the strict consensusthree nodes collapse (Fig. 27A). All trees support the mono-phyly of the classical Nephilinae (Clitaetra, (Herennia,(Nephilengys, Nephila))), excluding Phonognatha (contraHormiga et al. 1995) and Deliochus. Within the ingroup, theeight trees differ in the position of Clitaetra perroti andC. episinoides, which are either a grade or a clade sister to thethree African species. The sister clade to Clitaetra are theremaining nephilids, where the monophyly of Herennia,Nephilengys and Nephila are corroborated. The sister group ofnephilids (see below) is ambiguous (Fig. 27A) and is un-resolved in the strict consensus. Possible sister clades arearaneids, the clade with tetragnathids plus araneoid sheet-web builders, or the clade with all remaining araneoids.Deliochus is sister to Phonognatha, and the clade is sister to theremaining araneoids or alternatively sister to the group withtetragnathids and reduced piriform clade. Epeirotypus(symphytognathoids) and araneoid sheet web builders (Stea-toda, Nesticus, Linyphia, Pimoa) are monophyletic and sisterto tetragnathids, corroborating Hormiga et al. (1995) andGriswold et al. (1998). Tetragnathid monophyly excludingnephilines is also corroborated.

Successive weighting analysis stabilized after the seconditeration and resulted in a single tree identical to one of thefundamental cladograms (Fig. 27B). This working hypothesisof nephilid relationships hypothesizes Clitaetra perroti andC. episinoides as monophyletic and sister to the three Africanspecies, and nephilids as sister to the clade containing all

araneoids except Araneidae. Figure 28 shows unambiguouscharacter optimization on this phylogeny, and the alternative(ACCTRAN, DELTRAN) optimizations for the clitaetrineclade.

The phylogeny based on successive weighting (Fig. 27B)and the hypothesized character support (Fig. 28) form the basisfor the newly proposed nephilid classification and taxonomy.

TaxonomyFamily Nephilidae Simon, 1894 NEW RANK

According to the phylogenetic results, the group previouslyknown as Nephilinae with the genera Clitaetra, Herennia,Nephila and Nephilengys does not group with Tetragnathidaeand thus cannot continue to be catalogued there. Nephilidmonophyly, also corroborated by Kuntner (2005), and itsposition within Araneoidea will be retested within a largertaxonomic sample (Kuntner, Hormiga and Coddingtonunpublished) and its phylogenetic definition reassessed.

Phylogenetic definition. The family Nephilidae Simon, 1894is defined as the least inclusive clade, containing Clitaetra,Herennia, Nephila and Nephilengys.

Subfamily Clitaetrinae Simon, 1894

Monophyly. Four unambiguous synapomorphies are hypoth-esized for the clade Clitaetrinae (Fig. 28), identical incomposition to the genus Clitaetra (see Discussion): PLElarger than PME (15/1), sternal tubercle I absent (25/0), manyventromedian sclerotizations (58/1), and caudal copulatoryopening position (89/0). Further, ambiguous support(ACCTRAN optimization, Fig. 28) comes from: sternaltubercle III absent (27/0), female chillum absent (31/0), scle-rotized cymbial ectal margin (121/0), ventral tegular apoph-ysis (137/1), absence of embolic conductor membrane (145/0), finger-like embolic conductor shape (146/1), denticulatedembolus base (158/1), and side change on web (194/1).

Phylogenetic definition. The subfamily Clitaetrinae Simon,1894 and the genus Clitaetra Simon, 1889 are defined as theleast inclusive clade containing C. episinoides Simon, 1889(the type species of Clitaetra) and C. thisbe Simon, 1903. SeeDiscussion.

Genus Clitaetra Simon, 1889

Clitaetra Simon, 1889: 226. Type species by monotypy,Clitaetra episinoides Simon, 1889.Clitaetra: Bonnet 1956: 1100; Schmidt & Jocqué 1986: 211;Platnick 1989: 305; Platnick 1993: 371; Platnick 1997: 452;Dippenaar-Schoeman & Jocqué 1997: 292, 338. Platnick 2005.



Clytaetra: Simon 1894: 756; Roewer 1942: 925; Brignoli1983: 241. [Subsequent misspellings]Etymology. Unknown. The name is feminine in gender(Bonnet 1956: 1100).Monophyly. See Clitaetrinae, Figs 27, 28 and Discussion.Phylogenetic definition. Defined as Clitaetrinae, above (seeDiscussion).Diagnosis. Clitaetra differs from all non-nephilid spiders bythe striated cheliceral boss in both sexes (Figs 9C,D, 14A,Band 16A,B), and from all other nephilid genera by the pres-ence of a ventral tegular apophysis (Fig. 18A–D: VTA) andthe eye arrangement (Figs 1A, 3B, 5B, 6A, 7A, 9A, 12A, 16A,19A, 20A–C, 21A,B, 23A,B and 25A): lateral eyes widelyseparated, and posterior lateral eyes larger than the posteriormedians. It can further be distinguished from Nephila andNephilengys by the narrow cephalic region, dorsoventrallyflattened female abdomen and the sternum lacking sternalhumps, and from Herennia by the absence of abdominalsigillae, the absence of abdominal lobes, and by the smoothembolus without a hook.Note on conductor homology. Like other nephilids, Clitaetramales possess an extensive sclerite functioning as the conduc-tor, which entirely wraps the embolus (EC in Fig. 13).Schmidt & Jocqué (1986) referred to it simply as the embo-lus, but Hormiga et al. (1995) homologized it with the arane-oid conductor. While its homology with the conductor is stillone of the two alternative hypotheses, the sclerite is actuallywithin the embolic division (Kuntner 2005), and could betterbe described as the embolic conductor. This implies non-homology with the araneoid conductor, which is absent inClitaetra. The sclerite nomenclature here follows the term‘embolic conductor’ (EC).Description. Female (Figs 12, 14 and 15): Prosoma lengthfrom 1.6 to 3.5; total length from 3.5 to 9.9. Prosoma piri-form with narrow cephalic region. All eyes lack tapetum.Cheliceral boss with 50–100 striae (c. 300 in C. simoni), che-licerae with three prolateral and three or four retrolateralteeth and denticulated furrow (Fig. 3D). Sternum with threepaired groups of slit sensilla visible under dissecting micro-scope (Fig. 12C,D), numbers in each group varies from oneto five. Irregular patches of white pigment present onsternum. Legs with strong long spines on dorsal surface offemora, ventral, dorsal surface of tibiae. Opisthosoma oval topentagonal with three to five pairs of dorsomedian apodemesand no dorsolateral sclerotizations. Venter with 5–11 pairedmedian apodemes in rows, paired lateral row of sclerotiza-tions (Fig. 4A). Females (except C. thisbe) have two pairs ofwhite dots on posterior abdomen lateral to spinnerets, whichmay be fused into continuous white area (C. irenae). Epigy-num a simple sclerotized tongue (C. episinoides, C. perroti, seeFigs 12E,F and 19C,D), a ventral swelling (C. clathrata,C. simoni, C. irenae, Figs 1B–E, 6B,C and 7B,C), or a simple

transverse sclerotization (C. thisbe, Fig. 25B,C). Posteriorlyorientated small, inconspicuous copulatory openings, oftencovered by integument. Entelegyne vulva simple, pairedspermathecae connected to copulatory openings by tubularcopulatory duct (short, or coiled in C. irenae, Fig. 7D), ferti-lization duct originates adjacent. Spinnerets (Figs 4D–F,10E,F and 15C–F) of typical nephiline condition (see Hor-miga et al. 1995): ALS with ‘normal PI field’ where PI spigotbase is nearly as long or longer than the shaft (Griswold et al.1998: ch. 69, fig. 48B), major ampullate spigot and a nubbin,PMS with sparse aciniform field, and nubbin, PLS withaggregate spigots embracing flagelliform (observed inC. episinoides, C. clathrata, and C. irenae, condition unknownin C. simoni and C. thisbe) and with two cylindrical spigots ofnormal size, mesal being peripheral. Exception is C. perroti,where aggregate spigots of PLS triad were found apart fromflagelliforms (Fig. 22D–F).

Male (Figs 13A–D and 20A–G): prosoma length rangesfrom 1.3 to 2.5, total length from 2.8 to 4.9. Prosoma piri-form with narrow cephalic region. All eyes lack tapetum,lateral eyes widely separated, PLE larger than PME. Cheliceralboss striated (Fig. 16B), with fewer striae than in females.Chelicerae with three prolateral, three retrolateral teeth,5–15 cheliceral denticles. Sternum, legs as in female.Opisthosoma oval, dorsoventrally flattened, truncated poste-riorly (Fig. 20A–D), with dorsal scutum (Figs 17C–E and23E), three pairs of dorsal median apodemes. No dorsolateralsclerotizations. Ventral sclerotizations as in female (Figs 17Aand 23C). Two pairs of white dots on posterior abdomenlateral to spinnerets. Palp Nephila-like, with globular tegulumand prominent subtegulum (Fig. 13A) [in C. perroti subtegu-lum is distally fused to tegulum (Fig. 24A,C)]. Paracymbiumflat, rectangular, with invagination (Figs 5F, 13A, 18B and24B). Smooth narrow embolic conductor envelopes embolus,and is approximately at 90 degrees to bulb axis. Embolicconductor attached to embolus and tegulum via membrane(Fig. 13C), longer than cymbium, relatively straight(Fig. 13A–C). Embolus base separated from tegulum bymembrane (Fig. 20F). Embolus long, thin, positionedwithin embolic conductor groove (Fig. 13C). Embolussmooth, with no distal hooks or modifications. Prominentmacroseta present on distal part of palpal patella (Fig. 13A).Spinnerets without PLS triad in adults (Fig. 17F). Epian-drous gland spigot field sparse with 4–16 spigots (Figs 17Band 23D).Composition. The genus contains three clades (subgenera)with six species, one newly described.Natural history. Natural history is here presented for Clitae-tra irenae (see above), and some scattered information fromcollectors exists for C. episinoides and C. clathrata.Distribution. Western, central and southern Africa, Mada-gascar, Mayotte, Comoro Islands, Sri Lanka (Fig. 26).



Key to the Species of Clitaetra1(A) Female carapace edge and frontal opisthosoma withnumerous white setae (Fig. 9A,B), legs with dark brownarea around spine-sockets, epigynum a ventral swelling(Figs 1B–E, 6B,C and 7B,C). Embolic conductor wide(Figs 2A,B and 8A,B). African mainland species, subgenusAfroetra...................................................................................2(B) Female carapace edge and frontal opisthosoma withoutprominent setae, female legs unicolourous around spine-sockets, epigynum a tongue (Figs 12E,F and 19C,D) ora tranverse plate (Fig. 25B,C). Embolic conductor thin(Figs 13A–C and 20E,F). Species from Madagascar andIndian Ocean islands..............................................................42(A) Epigynum with distinct pair of copulatory openingsposteriorly (Fig. 7C), also visible from ventral view (Fig. 7B).Copulatory openings anterior to posterior sclerorized edge(Fig. 7C), long copulatory ducts coiling around fertilizationducts (Fig. 7D). Pale grey female prosoma surrounded by whitehairs, female dorsum white with lateral pair of red patches(Fig. 7A). Embolus fills almost half embolic conductor width(Fig. 8A). Southern African species............C. irenae sp. nov.(B) Epigynal copulatory openings not distinct and not seenfrom ventral view, positioned on posterior sclerorized edge(Figs 1B–E and 6B,C). Yellow-brown female prosoma,female abdomen with ‘butterfly’ pattern on anterior dorsum,and two pairs of dark patches on posterior dorsum (Figs 1Aand 6A). Embolic conductor broad, embolus filling roughlythird of it (Fig. 2A). Western African species.........................33(A) Females with grooved booklung covers, approximately150–250 boss striae (Fig. 3E), four retrolateral cheliceralteeth (Fig. 3D). Male palp as in Fig. 2A,B. Western Africa(Guinea-Bissau to Cameroon).................C. clathrata Simon(B) Females with smooth booklung covers, numerous bossstriae (c. 300), three retrolateral cheliceral teeth. Malesunknown. Central Africa (DR Congo)........C. simoni Benoit4(A) Epigynum a tongue with posteriorly hidden copulatoryopenings (Figs 12E,F and 19C,D). Spiders larger than 4.2 mm.Madagascar and adjacent islands, subgenus Clitaetra.............5(B) Epigynum a transverse plate with transparent area ante-riorly (Fig. 25B,C). Small spiders (3.5 mm). Sri Lanka, sub-genus Indoetra.................................................C. thisbe Simon5(A) Total female length 4.4–7.0. Female venter mediallywhite (Fig. 19B). Female carapace, abdomen broad (carapacelength to width ratio 1.1, abdomen length to width ratio 1.1).Epigynum a broad opaque tongue, spermathecae rarelyvisible through it (Fig. 19C,F). Spermathecae separated fromposterior tongue edge by one spermatheca diameter or less(Fig. 19E,F). Male subtegulum broken, distally fused totegulum (Fig. 20E,G and 24A,C). Embolus base distally withnumerous denticles (Figs 20F and 24D,E). Madagascar.......................................................................................C. perroti Simon(B) Total female length 4.9–9.0. Female venter medially dark

(Fig. 12C). Female carapace, abdomen elongated (carapacelength to width ratio 1.3, abdomen length to width ratio 1.4).Epigynum an elongated transparent tongue, spermathecaevisible through it (Fig. 12E). Spermathecae separated fromthe posterior tongue edge by two spermatheca diametersor more (Fig. 12E,G). Male subtegulum entire, separatedfrom tegulum (Figs 13A and 18A,B). Embolus base distallywith few denticles (Fig. 18D). Comoro Islands, Mayotte............................................................... C. episinoides Simon

Clade Afroetra subgen. nov.

Etymology. A combination of the words African and Clitaetra,the name is feminine in gender.Monophyly. Five unambiguous synapomorphies are hypo-thesized for the clade containing C. irenae, C. clathrata, andC. simoni (Fig. 28): female carapace edge with an extensive rowof hairs (4/1), dark spine socket colour (42/1), swollen epigy-nal ventral area (78/1), depressed epigynal anterior area (87/1), and curved epigynal lateral area (88/1). Further, ambigu-ous support (Fig. 28) comes from (ACCTRAN) groovedbooklung cover (75/0) and (DELTRAN) light and darkfemale sternum colour pattern (23/3).Phylogenetic definition. The subgenus Afroetra is defined asthe least inclusive clade containing C. clathrata Simon, 1889and C. irenae sp. nov.Diagnosis. Female carapace edge and frontal opisthosomawith numerous white setae (Fig. 9A,B), legs with dark brownarea around spine-sockets, epigynum a ventral swelling(Figs 1B–E, 6B,C and 7B,C). Embolic conductor wide(Figs 2A,B and 8A,B).Composition. The clade contains three species: C. clathrata,C. simoni and C. irenae sp. nov.Type species. Clitaetra irenae sp. nov.Distribution. Mainland Africa.

Clitaetra clathrata Simon, 1907 (Figs 1–5)

Clitaetra clathrata Simon, 1907: 283–284, description offemale (from Guinea Bissau). Female holotype without local-ity label in MCSNG, label reading ‘Clytaetra clatrata Simon’[note misspellings of both the generic and the specific name],examined. Simon (1907) cites the locality as ‘Guinée portug.:Rio Cassine’. Approximate coordinates for Rio Cassine(GUINEA BISSAU) are 10°18′N 15°24′W.Clytaetra clathrata: Roewer 1942: 925.Clitaetra clathrata: Bonnet 1956: 1100; Benoit 1962: 221–222,description of female; Platnick 2005.Clitaetra sp.: Hormiga, Eberhard & Coddington 1995:fig. 9A–C, male palpal anatomy.Additional material examined. CAMEROON: South-westProv., Fako Division, Limbe Subdiv., south slope of Mt.



Etinde, nr. Batoke, 4°3′N, 9°6′E, 300–1000 m, 10 January1992 (coll. Griswold et al.) — CL019/j1 (Immature), CL019/m1 (Male); 1.4 km NE of Etome, 4°2.58′N, 9°7.31′E, 400,13–19 January 1992 (coll. Larcher et al.) — CL001/j1 (Imma-ture), cl017/j1 (Immature), cl001/m1 (Male), cl001/m2(Male), cl017/m1 (Male), cl017/m2 (Male), cl017/m3 (Male),CL018/m1 (Male); COTE D’IVOIRE: Appouesso, FCBossematie, route nr 1, 6°36′N, 3°27′W, rain forest, layon 19;in centre of web together with cocoon, 19 February 1997(coll. R.Jocque & L.Baert) — cl014/f1 (Female), cl014/j1(Immature), cl014/m1 (Male).Etymology. Unknown.Diagnosis. Clitaetra clathrata females differ fromC. episinoides, C. perroti, and C. thisbe by the carapace andfrontal abdomen with numerous white setae, by the darkbrown area around leg spine-sockets, and by the female epigy-num with a ventral swelling (Fig. 1B–E). Clitaetra clathratafemales differ from C. irenae by having the copulatory open-ings positioned on the posterior sclerotization (Fig. 1C,E).Clitaetra clathrata females can be distinguished from C. simoniby having grooved booklung covers, fewer cheliceral bossstriae (c. 150–250, Fig. 3E), by four retrolateral cheliceralteeth (Fig. 3D), and by the posterior epigynum lacking anotched appearance (Fig. 1C,E). A unique feature of

C. clathrata is the thorny abdominal cuticle of the female(Fig. 4C). Male C. clathrata can be distinguished from those ofall other species by the short and wide embolic conductor: theembolus width (in mesal view) is less than one third of themaximum length of the embolic conductor (Fig. 2B).Description. Female (cl14/f1, from Ivory Coast, compare withFigs 1 and 3,4): Total length 7.4. Prosoma (Fig. 1A) 3.4 long,2.8 wide, 2.1 high; yellow-brown. Cephalothorax, frontalabdomen surrounded by numerous white setae (Fig. 3B).Sternum 1.4 long, 1.4 wide; medially white, laterally brown,with irregular patches of white pigment. AME diameter 0.17,ALE 0.13, PME 0.14, PLE 0.16. AME separation 0.20, PMEseparation 0.28, PME–PLE separation 0.22, AME–ALEseparation 0.08, AME–PME separation 0.32, ALE–PLE separa-tion 0.28. Both eye rows recurved (Figs 1A and 3B). Clypeusheight 0.16. Chelicerae (Fig. 3C,D) with three prolateral andfour retrolateral teeth (the fourth smaller and positionedslightly medially), with approximately 15 cheliceral denticles(see Variation). Cheliceral boss with roughly 200 striae (seeVariation). Appendages. Legs yellow-light brown, with femora,patellae, tibiae, metatarsi and tarsi distally darker brown;dark brown pigment spots around macrosetae bases. Leg Ilength 10.9 (Fe 5.0, Pa 1.0, Ti 3.9, Me 5.3, Ta 1.7). Legformula 1-2-4-3. Opisthosoma 4.7 long, 3.4 wide, 2.0 high,

Fig. 1 —A–E. Clitaetra clathrata, femalemorphology: —A. Habitus, dorsal, fromGuinea-Bissau (holotype). —B. Epigynum,ventral (same). —C. Epigynum, posterior(same). —D. Epigynum, ventral, from IvoryCoast (cl14/f1). —E. Epigynum, posterior(same). — Scale bars: A = 1.0 mm, B–E =0.1 mm.



pentagonal, wider posteriorly, with four pairs of dorsalapodemes, as in Fig. 1A. Dorsum pattern as in Fig. 1A withwhite lines demarcating four darker areas posteriorly, andwith dark brown anterior butterfly-shaped patch. Venter withpatches of white and grey, medially whitish, caudally with twopairs of white dots. Eight pairs of ventromedian apodemes,and paired row of ventrolateral sclerotizations (Fig. 4A,B),the latter not clearly discernible with a dissecting micro-scope. Epigynum as in Fig. 1D,E (Ivory Coast) and Fig. 1B,C(holotype). Epigynum plate (Fig. 1B–E) has ventral swellingwith paired lateral curve, anterior depression, copulatoryopenings on posterior sclerotized edge.

Male (cl18/m1, from Etome, Cameroon, Figs 2 and 5):total length 3.9. Prosoma (Fig. 5B) 1.8 long, 1.6 wide, 1.1high; red-brown. Sternum 0.9 long, 0.8 wide; orange-brown.AME diameter 0.16, ALE 0.11, PME 0.09, PLE 0.11. AMEseparation 0.12, PME separation 0.19, PME–PLE separation0.14, AME–ALE separation 0.03, AME–PME separation0.20, ALE–PLE separation 0.17. Both eye rows recurved(Fig. 5B). Clypeus height 0.11. Chelicerae red-brown withthree prolateral, three retrolateral teeth, five denticles. Che-liceral boss with about 40 striae. Appendages. Legs orange-brown; tibiae, metatarsi distally darker. Leg I length 10.6 (Fe2.8, Pa 0.6, Ti 2.5, Me 3.13, Ta 1.33). Leg formula 1-2-4-3.Opisthosoma 2.5 long, 1.3 wide, 1.0 high, oval, with lightbrown scutum, four pairs of dorsal apodemes. Ventrally grey,

caudally with two pairs of white dots around spinnerets.Ventro-median apodemes, ventrolateral sclerotizations as infemale. Pedipalp (Figs 2A–C and 5B–F) with suboval tegulum,prominent, separate subtegulum, prominent ventral apophy-sis. Embolic conductor short, wide, enveloping embolus.Embolus base distally with few denticles (Fig. 5E).Variation. Females: prosoma length ranges from 2.6 to 3.4(n = 5); total length from 5.4 to 7.4 (n = 5). Prosoma and legsvary in coloration from light yellow to dark brown. Prosomamay exhibit two dark lateral and a median band (not in holo-type). Irregular patches of white pigment present on sternumof most preserved females. Number of cheliceral denticles15–38. Cheliceral boss with 150–250 striae. Leg formula 1-2-4-3 (females from Ivory Coast) or 1 = 2, 4, 3 (holotype).Dorsum pattern varies in contrast and coloration, but gener-ally as illustrated (Fig. 1A). Venter pattern variable but medi-ally lighter. Seven or eight pairs of ventromedian apodemes.Spermathecae visible through cuticle in most females.

Males: prosoma length ranges from 1.5 to 1.9 (n = 7), totallength from 2.8 to 3.9 (n = 7). Ejaculatory duct with single(Fig. 2C, from Cameroon) or double switchback in tegulum(Fig. 2D, from Ivory Coast). Fourth (small) pair of dorsalapodemes may be absent. Prosoma, leg coloration from lightyellow to orange-brown. Dorsum uniform grey to brown, orwith darker grey midline band in front, which continues intoposterior dark part across abdomen width.

Fig. 2 —A–D. Clitaetra clathrata, male leftpedipalp: —A. Ectal, from Cameroon(cl18/m1). —B. Mesal (same). —C. Bulbtransparent, showing sperm duct, fromCameroon (cl17/m1). —D. Same, fromIvory Coast (cl14/m1). Scale bars = 0.1 mm.



Distribution. Western Africa, from Guinea Bissau toCameroon (Fig. 26).Natural history. Label data suggest that the species inhabitsrain forests of western Africa of elevations up to 1000 m, andthe female builds an orb web with egg sac suspended in it.

Clitaetra simoni Benoit, 1962 (Fig. 6)

Clitaetra simoni Benoit, 1962: 222, description of female(from DR Congo). Female holotype in RMCA (M.T. 12467):‘Congo. Sankuru: Komi, iv-1930 ( J. Ghesquière)’, examined.The type locality lies approximately at 03°23′S, 23°46′E.Clytaetra simoni: Brignoli, 1983: 241.Clitaetra simoni: Platnick, 2005.Additional material examined. None.

Fig. 3 A–F. Clitaetra clathrata, female prosoma, from Ivory Coast (cl14): —A. Frontal. —B. Dorsal. —C. Chelicerae, ventroectal. —D. Rightchelicera, ventromesal. —E. Right chelicera, ectal, with ridged cheliceral boss (arrow). —F. Sternum, with numerous slit sensilla (arrows). Scalebars: A–C = 100 µm, D–F = 10 µm.



Note. The species diagnosis below separates the holotypefemale of C. simoni from morphologically similar C. clathrata.In the absence of available males and additional female mate-rial, I here tentatively recognize C. simoni as valid.Etymology. The species epithet was devised as a patronymhonouring Eugene Simon (see Taxonomic History).Diagnosis. Clitaetra simoni female differs from C. episinoides,C. perroti, and C. thisbe by the carapace and frontal abdomen

with numerous white setae, by the dark brown area aroundleg spine-sockets, and by the female epigynum with a ventralswelling (Fig. 6B,C). Clitaetra simoni female differs from thatof C. irenae by having the copulatory openings positioned onthe posterior sclerotization (Fig. 6C). The species resemblesC. clathrata in general somatic morphology, but can be distin-guished from the latter by the smooth booklung covers, morenumerous boss striae (c. 300), and by the three retrolateral

Fig. 4 A–F. Clitaetra clathrata, female opisthosoma, from Ivory Coast (cl14): —A. Venter, with paired rows of sclerotizations (arrows). —B.Ventrolateral sclerotizations. —C. Detail of (lateral) opisthosoma. —D. Spinnerets. —E. Anterior lateral spinnerets. —F. Posterior spinnerets.— Scale bars: A, B, D = 100 µm, C, E, F = 10 µm.



cheliceral teeth. The epigynum (Fig. 6B,C) has small andunsclerotized anterior depressions and sclerotized edgesof the lateral curve. Spermathecae are not visible throughthe epigynal plate, as they are in C. clathrata. The posteriorepigynum has a notched appearance (Fig. 6C).Description. Female (holotype): Total length 8.7. Prosoma(Fig. 6A) 3.4 long, 3.1 wide, 2.2 high; yellow-brown. Carapaceedge, frontal abdomen with numerous white setae. Ster-

num 1.4 long, 1.5 wide; medially white, laterally dark, withthree paired groups of slit sensilla. AME diameter 0.18, ALE0.16, PME 0.14, PLE 0.15. AME separation 0.23, PME sep-aration 0.35, PME–PLE separation 0.23, AME–ALE separa-tion 0.12, AME–PME separation 0.35, ALE–PLE separation0.26. Both eye rows recurved (Fig. 6A). Clypeus height 0.13.Chelicerae with three prolateral and three retrolateral teeth, andapproximately 28 denticles. Cheliceral boss with approximately

Fig. 5 A–F. Clitaetra clathrata, male, from Cameroon (cl17/m3): A–C. Prosoma. —A. Frontal. —B. Dorsal. —C. Ventral. —D–F. Left pedipalp.—D. Dorsoectal. —E. Mesal. —F. Ectal, detail of paracymbium, with a folding of the margin (arrow). — Scale bars: A–E = 100 µm, F = 10 µm.



300 striae. Appendages. Legs yellow. Femur I length 4.6 (otherjoints lost). Leg formula unknown due to poor condition ofholotype. Opisthosoma 6.5 long, 4.9 wide, 3.9 high. Opistho-soma as in C. clathrata, but more oval and less pentagonal,dorsal pattern less pronounced, blurred (Fig. 6A). Ventrally asin C. clathrata, caudally with two pairs of white dots. Eightpairs of ventromedian apodemes, two posterior pairs ofventrolateral sclerotizations. Epigynum (Fig. 6B,C) a ventralswelling with posterior notched appearance.

Male: unknown.Distribution. Known only from the type locality, DemocraticRepublic of the Congo (Fig. 26).Natural history. Unknown.

Clitaetra irenae sp. nov. (Figs 7–11)

Clitaetra sp.: Hormiga, Eberhard & Coddington 1995:fig. 15A–D, female spinneret anatomy.Types. Female holotype (CL30, in PPRI), two male paratypes(CL29 and CL31, in USNM) from SOUTH AFRICA,KwaZulu-Natal, St. Lucia Estuary, Fanies Island, 28°6′41S,32°25′51′′E, Kuntner, Agnarsson, Hormiga, Coddington, 31March–5 April 2001. Two female paratypes (CL2, in CAS)from (locality from museum label): South Africa, Natal, St.Lucia National Park, Fanies Camp, 28°S, 32°30′E, V. & B.Roth, 24 January 1991.Additional material examined. MALAWI: [Nyasaland], 53 mi.N of Blantyre, 15°5′S, 34°59′E, 630, 25 February 1958 (coll.Ross & Leech) — cl047/m1 (Male); SOUTH AFRICA:

Kwazulu Natal, Sodwana Bay (area), 27°30′S, 32°30′E, orbweb against tree trunks in Dune Forest, 25 September 1998(coll. A. Leroy) — CL020/f1 (Female); KwaZulu-Natal, St.Lucia Estuary, Fanies Island, 28°6.41′S, 32°25.51′E, 25, orbwebs against tree trunks, 31 March−5 April 2001 (coll. Kunt-ner et al.) — cl030/f1 (Female), cl032/f1 (Female), cl034/f1(Female), cl035/f1 (Female), cl044/f1 (Female), cl029/j1(Immature), cl033/j1 (Immature), cl044/j1 (Immature),cl029/m1 (Male), cl031/m1 (Male), cl034/m1 (Male), cl035/m1 (Male), cl044/m1 (Male); Makakatana Bay, 28°14.13′S,32°24.38′E, 25, on a wooden wall of a latrine, 1 April 2001(coll. Kuntner et al.) — cl027/f1 (Female); Phinda ResourceReserve, South sand forest, 27°50.43′S, 32°18.49′E, 13 April2001 (coll. Kuntner et al.) — cl041/f1 (Female); NtabankosiMt. rock outcrops, 27°51.52′S, 32°16.3′E, 14 April 2001(coll. Kuntner et al.) — cl043/j1 (Immature), cl043/m1(Male); North sand forest, 27°46.21′S, 32°19.56′E, 16 April2001 (coll. Kuntner & Agnarsson) — cl042/j1 (Immature);Sodwana Bay, Mgoboseleni Trail (and Sodwana camp area),27°32.0′S, 32°39.5′E, 10, 6–7 April 2001 (coll. Kuntner et al.)— cl036/f1 (Female), cl037/f1 (Female), cl036/j1 (Imma-ture); Maputuland, Tembe Elephant Park, 27°2′S, 32°25′E,Orb web on tree trunk with lower half enlarged, 10 April1998 (coll. J Leeming) — CL021/f1 (Female); Tembe Ele-phant Park, Ngobozana sand forest, 27°2.51′S, 32°25.19′E,100, 9–12 April 2001 (coll. Kuntner et al.) — cl026/f1(Female), cl028/f1 (Female), cl026/j1 (Immature), cl028/m1(Male); (coll. Kuntner et al.) — cl038/f1 (Female), cl039/f1(Female), cl039/j1 (Immature), cl040/j1 (Immature), cl040/

Fig. 6 A–C. Clitaetra simoni, female holo-type from DR Congo: —A. Habitus, dorsal.—B. Epigynum, ventral. —C. Epigynum,posterior. Scale bars: A = 1.0 mm, B, C = 0.5 mm.



m1 (Male); St. Lucia National Park, Fanies Camp, 28°0′S,32°30′E, 24 Jan 1991 (coll. V. D. & B. Roth) — CL002/f1(Female).Etymology. The species epithet, a noun in genitive case, is apatronym honouring my wife Irena Kuntner.Diagnosis. Copulatory openings of C. irenae females are noton the posterior sclerotized epigynal margin, as in all otherClitaetra species, but are well anterior to the margin(Fig. 7C), within a pair of sclerotizations. The shape of thelong copulatory ducts coiling around the fertilization ducts(Fig. 7D) is unique within the genus. Males can be distin-guished from other Clitaetra species by the shape of the tegu-lum (broader than long, Fig. 8A) and the embolic conductor(Fig. 8A,B), which is relatively longer than in C. clathrata andstraighter than in C. episinoides.Description. Female (paratype CL2f1 from Fanies Camp,CAS): Total length 7.1. Prosoma (Fig. 7A) 3.2 long, 2.6wide, 1.7 high; prosoma, chelicerae yellow-grey. Carapaceedge, frontal abdomen with numerous white setae (Fig. 9A,B).Sternum 1.34 long, 1.26 wide, yellow, with irregular whitepatches along middle, with four pairs of sternal slit sensilla(Fig. 9E,F). AME diameter 0.17, ALE 0.11, PME 0.11, PLE0.13. AME separation 0.19, PME separation 0.25, PME–PLE separation 0.19, AME–ALE separation 0.08, AME–PME separation 0.26, ALE–PLE separation 0.24. Clypeusheight 0.16. Chelicerae with three prolateral and three retro-

lateral teeth, and 24 denticles. Cheliceral boss not prominent,striae hardly visible with dissecting scope (but see variationand Fig. 9C,D). Appendages. Legs whitish-yellow, with darkbrown spine-sockets and spines; tarsi brown. Sustentaculum(tarsus IV) conspicuous. Leg I length 15.3 (Fe 4.5, Pa 1.3, Ti3.6, Me 4.3, Ta 1.7). Leg formula 1-2-4-3. Opisthosoma(Figs 7A and 10A) 4.4 long, 3.3 wide, 2.3 high, pentagonal,dorsoventrally flattened, with five pairs of dorsomedian apo-demes (fourth and fifth inconspicuous); dorsum with grey,brown, white pattern (illustrated in Fig. 7A from holotype),venter with broad white midband extending from epigastricfold to spinnerets, and broken white pattern around spin-nerets. Eleven pairs of ventromedian apodemes, two posteriorpairs of inconspicuous ventrolateral sclerotizations visibleunder dissecting microscope (but, paired row of lateral scle-rotizations visible on SEM images, Fig. 10A,B). Epigynum(illustrated from paratype CL2f2) as diagnosed (Figs 7B–Dand 10C,D).

Male (paratype CL29m1 from Fanies Island, USNM):Total length 3.1. Prosoma 1.7 long, 1.3 wide, 0.8 high; lightbrown. Sternum 0.70 long, 0.73 wide; light brown. AMEdiameter 0.13, ALE 0.06, PME 0.06, PLE 0.07. AME separa-tion 0.12, PME separation 0.18, PME–PLE separation 0.11,AME–ALE separation 0.03, AME–PME separation 0.18,ALE–PLE separation 0.16. Both eye rows recurved. Clypeusheight 0.1. Appendages. Legs orange-brown. Leg I length 9.6

Fig. 7 A–D. Clitaetra irenae sp. nov., femalemorphology, from South Africa: —A. Habitus,dorsal (holotype). —B. Epigynum, ventral(paratype cl2/f2). —C. Epigynum, posterior(same). —D. Epigynum, cleared, dorsoectal(same). Scale bars: A = 1.0 mm, B–D = 0.1 mm.



(Fe 2.5, Pa 0.6, Ti 2.1, Me 2.6, Ta 1.2). Leg formula 1-2-4-3. Opisthosoma 1.9 long, 1.1 wide, 0.6 high, oval, with threepairs of dorsal apodemes (Fig. 11A,B). Scutum dark brown,with white frontal band. Venter brown, dark grey, caudallywith three pairs of white dots around spinnerets. Pedipalp asin Figs 8 and 11C–F. Tegulum suboval (broader than long),with prominent ventral apophysis (Figs 8A–C and 11C–E).Embolic conductor long, wide, embolus filling roughly halfits width (Fig. 8A,B).Variation. Females: total length ranges from 5.7 to 9.9(n = 12), prosoma length ranges from 2.7 to 3.5 (n = 12). Littlepigmented prosoma and legs can be white or pale yellow-brown. In most females a pair of dark spots is present on thecarapace anterior to the depression. Chelicerae with three orfour retromarginal teeth, 11–24 denticles. 100–200 (Fig. 9D)cheliceral boss striae difficult to observe due to lack ofpigment. Distal parts of leg joints in some specimens darkbrown. In some females white setae on carapace and abdo-men edge are so abundant to give the animal velvety appear-ance. Most females (holotype in Fig. 7A, not paratype CL2f1

described) with pair of red patches laterally on ornatedorsum. Booklung covers of some exemplars with grooves.Some females have a truncated posterior abdomen. 8–11pairs of ventromedian apodemes. In some females copulatoryopenings are not visible without removing posterior tissue(Fig. 10D). Some copulatory openings covered with hard redmatter.

Males: Total length ranges from 2.8 to 3.6 (n = 4), prosomalength from 1.4 to 1.9 (n = 4). Coloration is grey to orange.Distribution. South-east Africa: South Africa (KwaZulu-Natal), Malawi (Fig. 26).Natural history. See Biology.

Clade Clitaetra s.s.

Etymology. The nominal subgenus, see above.Monophyly. A single unambiguous synapomorphy is hypo-thesized for the clade containing C. episinoides and C. perroti(Fig. 28): tongue shaped epigynal posterior area (79/1). Fur-ther, ambiguous support (Fig. 28) comes from (DELTRAN)

Fig. 8 A–C. Clitaetra irenae sp. nov., male leftpedipalp, from South Africa (paratype cl29/m1): —A. Ectal. —B. Mesal. —C. Bulbtransparent, showing sperm duct, ectal. Scalebars = 0.1 mm.



wider than long female sternum (21/1), smooth booklungcover (75/1), denticulated embolus base distal part (158/1).Phylogenetic definition. The subgenus Clitaetra is defined asthe least inclusive clade containing C. episinoides Simon, 1889and C. perroti Simon, 1894.Diagnosis. Epigynum tongue-shaped with posteriorly hidden

copulatory openings (Figs 12E,F and 19C,D). Females largerthan 4.2 mm.Composition. The clade contains two species, C. episinoidesand C. perroti.Type species. Clitaetra episinoides Simon.Distribution. Madagascar and adjacent islands (Fig. 26).

Fig. 9 A–F. Clitaetra irenae sp. nov., female prosoma, from South Africa (cl28/f1): —A. Dorsolateral, box defines area of image B. —B. Carapaceedge. —C. Chelicerae and head region, ventroectal, box defines area of image D. —D. left cheliceral boss, ventroectal. —E. Ventral, box definesarea of image F. —F. Detail of sternum, with slit sensilla (arrows). Scale bars = 100 µm.



Clitaetra episinoides Simon, 1889 (Figs 12–18)

Clitaetra episinoides Simon, 1889: 226–227, description offemale (from Mayotte). Two syntype females in MNHN,labelled ‘4979 Clit. episinoides E. S. Mayotte’, examined.The type locality, Mayotte, lies approximately at 12°50′S,45°10′E.

Clytaetra episinoides: Simon 1894: 756, f.834; Petrunkevitch1928: 142; Roewer 1942: 925.Clitaetra episinoides: Simon, 1907: 284; Bonnet, 1956: 1100;Schmidt & Jocqué, 1986: 211, f.6, description of male; Plat-nick, 1989: 305; Platnick, 1997: 452; Platnick, 2005.Additional material examined. COMOROS: Grande Comore(Njazidja), Boboni, cave, 11°44′S, 43°17′E, 600, in a cave,

Fig. 10 A–F. Clitaetra irenae sp. nov., female opisthosoma, from South Africa (cl28/f1): —A. Venter, with paired rows of sclerotizations (arrows).—B. Detail of venter with ventrolateral sclerotizations (arrows). —C. Epigynum, ventral. —D. Epigynum, posterior, with copulatory openings(arrow). —E. Spinnerets. —F. lateral spinnerets. Scale bars: A, C, D = 100 µm, B, E, F = 10 µm.



27 November 1983 (coll. Jocqué) — cl006/f1 (Female), cl006/j1(Immature), cl006/m1 (Male), cl006/m2 (Male); M’Vouni,11°35′S, 43°20′E, 440, forest, 6 August 1981 (coll. Jocqué) —cl007/m1 (Male); M’Vouni, 11°35′S, 43°20′E, 400–700,8 December 1983 (coll. Jocqué) — cl008/m1 (Male); [FRANCE]:Mayotte, Coconi, DAF Campus, 12°47′S, 45°9′E, 15 February1999 (coll. Jocqué) — cl010/f1 (Female); Convalescence, ruins,

12°47′S, 45°13′E, ruins, 16 February 1999 (coll. Jocqué & DeSmet) — cl009/f1 (Female); Mayotte, 12°48′S, 45°9′E—cl023/f1 (Female); Mt. Benara, 12°52.0′S, 45°10.36′E, 230,sweeping along track to reserve gate, 23 February 1999 (coll.R. Jocque & DeSmet) — cl011/f1 (Female).Etymology. The species epithet reflects the ‘appearance and colora-tion of an Episinus’ (Simon 1894: 756), a theridiid spider genus.

Fig. 11 A–F. Clitaetra irenae sp. nov., male, from South Africa (cl28/m1): —A. Lateral opisthosoma with a dorsal scutum (arrow). —B. Detailof scutum with an apodeme (arrow). —C–F. Right pedipalp. —C. Ectal. —D. Bulb, ectal. —E. Apical. —F. Embolic conductor tip, apicallybroken. Scale bars = 10 µm, except C = 100 µm.



Diagnosis. The females of C. episinoides can be distinguishedfrom all other Clitaetra species except C. perroti by the pres-ence of a ‘tongue’ epigynum (Figs 12E,F and 15B) withposteriorly hidden copulatory openings. Clitaetra episinoidesdiffers from C. perroti by the combination of the followingcharacters: epigynum an elongated, well sclerotized buttransparent tongue with spermathecae visible anterior to thetongue (Fig. 12E); spermathecae separated from the poste-rior epigynum tongue edge by two spermatheca diameters ormore (Fig. 12E,G); female venter medially dark (Fig. 12C);female carapace elongated (carapace length to width ratio1.3); opisthosoma long (length to width ratio 1.4). MaleC. episinoides differ from C. irenae and C. clathrata by thenarrow and curved embolic conductor (Fig. 13A,B). Unlikein C. perroti, the subtegulum of C. episinoides males is not fusedto the tegulum (Figs 13A and 18A,B), and the embolus base has

few denticles distally (Fig. 18D). The ratio of embolic con-ductor length to cymbium length in male palp when observedfrom ectal side is more than 1.5 : 1 and the sperm duct in thetegulum (Fig. 13A,D) is relatively narrower than in C. perroti.Description. Female (cl9/f1, from Convalescence, Mayotte;Figs 12 and 14,15): Total length 7.9. Prosoma (Fig. 12B) 3.3long, 2.5 wide, 2.0 high; light brown. Chelicerae dark brown.Sternum 1.3 long, 1.3 wide, with small paired tubercle adja-cent to coxa 4 (Fig. 14D) and three paired groups of slitsensilla (Fig. 12D); dark brown laterally, medially light, withirregular white pigment patches, labium dark brown. AMEdiameter 0.19, ALE 0.14, PME 0.14, PLE 0.19. AMEseparation 0.19, PME separation 0.25, PME–PLE separation0.2, AME–ALE separation 0.1, AME–PME separation0.25, ALE–PLE separation 0.19. Both eye rows recurved(Fig. 12B). Clypeus height 0.28. Chelicerae with three

Fig. 12 A–G. Clitaetra episinoides, femalemorphology, from Mayotte (cl9/f1–3):—A. Habitus, lateral. —B. Habitus, dorsal.—C. Habitus, ventral. —D. Sternum(note slit sensilla). —E. Epigynum,ventral. —F. Epigynum, posterior. —G.Epigynum, cleared, dorsal (leftspermatheca displaced). Scale bars: A–C = 1.0 mm, D–F = 0.5 mm, G = 0.1 mm.



prolateral, three retrolateral teeth, approximately 22 den-ticles (Fig. 14C). Cheliceral boss with roughly 130 striae(Fig. 14B). Appendages. Legs yellow-light brown, with darkbrown rings, notably on femora, tibiae. Leg I length 21.0 (Fe6.0, Pa 1.2, Ti 5.5, Me 6.2, Ta 1.9). Leg formula 1-2-4-3.Opisthosoma (Fig. 12A–C) 4.8 long, 3.2 wide, 2.1 high, widestposteriorly, with three conspicuous pairs of dorsal apodemes,fourth inconspicuous. Dorsum with white, grey, brownpattern as in Fig. 12B, venter medially dark, delimited by pairof mid-lateral white lines (Fig. 12C). Epigynum a prominent,sclerotized tongue, with spermathecae visible through it(Figs 12E and 15B), inconspicuous copulatory openings onposterior sclerotization (Fig. 12F). Inner epigynum simple(Fig. 12G), with spermathecae connected to copulatoryopening by short, tubular copulatory duct.

Male (cl6/m1, from Boboni, Comoro Islands;Figs 13,16,17,18): Total length 4.9. Prosoma 2.5 long, 1.9

wide, 1.4 high; light brown. Sternum 1.13 long, 1.08 wide;light brown. AME diameter 0.2, ALE 0.13, PME 0.11, PLE0.14. AME separation 0.13, PME separation 0.19, PME–PLE separation 0.20, AME–ALE separation 0.04, AME–PME separation 0.24, ALE–PLE separation 0.19. Both eyerows recurved. Clypeus height 0.16. Chelicerae light brown,with three prolateral, three retrolateral teeth, approximately15 denticles. Cheliceral boss (Fig. 16A,B) with roughly 100striae. Appendages. Legs yellow-light brown, with dark brownrings, notably on femora, tibiae. Leg I length 21.8 (Fe 5.9, Pa1.1, Ti 6.0, Me 6.7, Ta 2.1). Leg formula 1-2-4-3. Opisthosoma(Fig. 17) 2.9 long, 1.6 wide, 1.2 high, oval, dorsoventrallyflattened, truncated posteriorly, dorsally with three pairs ofapodemes, ventrally with paired row of median apodemes,lateral sclerotizations (Fig. 17A). Dorsum with white dots,darker midband, venter grey, with black midband. Pedipalp asin Figs 13 and 18: globular tegulum, prominent, separate

Fig. 13 A–D. Clitaetra episinoides, malepedipalp, from Comoro Islands: —A.Left pedipalp, ectal (cl6/m2). —B.Same, mesal. —C. Right pedipalp,expanded (cl6/m2). —D. Left bulb,transparent, showing sperm duct, ectal(cl8/m1). Scale bars = 0.1 mm.



subtegulum, long, narrow, slightly curved embolic conductorenveloping embolus. Embolus long, thin, smooth (Fig. 13C).Variation. Females: prosoma length ranges from 2.3 to 3.4(n = 8); total length from 4.9 to 9.0 (n = 8). Prosoma and legsvary in coloration in preserved specimens from pale greyto yellow. Irregular patches of white pigment are present onsternum of most preserved females. Cheliceral denticle

numbers 12–27. Cheliceral boss striae 60–150. Dorsum andventer patterns vary in contrast and coloration. Seven oreight pairs of ventromedian apodemes are present. Pairedrow of ventrolateral sclerotizations is present but difficult toobserve under light microscope.

Males: prosoma length ranges from 2.0 to 2.5 (n = 4); totallength from 4.1 to 4.9 (n = 4). Prosoma and legs vary in

Fig. 14 A–F. Clitaetra episinoides, female, from Mayotte (cl9): —A. Prosoma, ectal, box defines area of image B. —B. Left chelicera, ectal, withridged cheliceral boss (arrow). —C. Chelicerae and maxillae, ventroectal. —D. Sternum with paired tubercle (arrow) adjacent to coxae 4,ventral. —E. Detail of patella 1 with a paired lyriform organ (arrows), prolateral. —F. Tarsus 4 showing the claws and sustentaculum (arrow).— Scale bars: A, C, D = 100 µm, B, E, F = 20 µm.



coloration in preserved specimens from yellow to red-brown.Scutum pronounced in most males, dorsum colour variable.Cheliceral denticle numbers 11–15. Ventral apodemes andsclerotizations as in female.Distribution. Known only from the Indian Ocean islandsMayotte (Fr.) and Njazidja (= Grande Comore, ComoroIslands) (Fig. 26). Contra Hormiga et al. (1995) and Platnick(2005) the species does not occur in Madagascar (see below).

Note on distribution. There has been confusion about thegeographic distribution of Clitaetra episinoides. Simon (1889)described it from the French island of Mayotte in the IndianOcean. When describing the second species of the genus,C. perroti from Madagascar, Simon (1894) provided readerswith a generic diagnosis, listing its type species (C. episinoides),and the distribution of the genus (islands Mayotte andMadagascar). Roewer (1942: 925) listed the distribution of

Fig. 15 A–F. Clitaetra episinoides, female opisthosoma, from Mayotte (cl9): —A. Detail of dorsal abdomen with apodeme. —B. Epigynum,ventral. —C. Spinnerets. —D. ALS. —E. PMS. —F. PLS. Scale bars = 20 µm, except B = 100 µm.



C. episinoides as ‘Mayotta’, but Bonnet (1956: 1100) errone-ously cited the distribution as ‘Madagascar, Comores’. Thesame is true for the catalogues of Platnick (1989: 305, 1997:452, 2005). The species does not occur on Madagascar.Natural history. See ‘Biology’ of Clitaetra. Label data fromGrande Comore suggest the species can be found at eleva-tions up to 700 m.

Clitaetra perroti Simon, 1894 (Figs 19–24)

Clytaetra Perroti Simon, 1894: 756–757, description of female(from Madagascar). Female holotype in MNHN, labelled‘10224 Clit. Perroti E. S. Tamatave’ [= Toamasina, approxi-mately at 18°8′S, 49°24′E], examined.Clitaetra Perroti: Simon 1907: 284.

Fig. 16 A–F. Clitaetra episinoides, male prosoma, from Comoro Islands (cl6/m2): —A. Ectal, box defines area of image B. —B. Cheliceral boss.—C. Ventral. —D. Ventroectal. —E. Dorsal, box defines area of image F. —F. Detail of posterior carapace. Scale bars = 100 µm, except B = 10 µm.



Clytaetra perroti: Roewer 1942: 925.Clitaetra perroti: Bonnet 1956: 1100; Platnick (2005).Clitaetra episinoides: Hormiga, Eberhard & Coddington 1995:330, f. 9D,E (male), misidentification.Additional material examined. MADAGASCAR: Dist. Fian-aratsoa, Ambozotany, 21°15′S, 47°19′E, introduced pineforest, 15 July 1992 (coll. V. & B.Roth) — cl015/f1 (Female),

cl015/f2 (Female), cl015/f3 (Female), cl015/f4 (Female),cl015/f5 (Female), cl015/f6 (Female), cl015/f7 (Female),cl015/j1 (Immature), cl015/m1 (Male), cl015/m2 (Male),cl015/m3 (Male); Ranomafana N.P., Ambozotany, 21°25′S,47°33′E, 4 June 1992 (coll. Barbara Roth) — cl016/f1(Female), cl016/m1 (Male); Foulpointe, 17°40′S, 49°31′E,forest on sand, Pandanus swamp, November 1993 (coll.

Fig. 17 A–F. Clitaetra episinoides, male opisthosoma, from Comoro Islands (cl6/m2): —A. Venter, with paired rows of sclerotizations (arrows),box defines area of image B. —B. Epiandrous gland spigots. —C. Posterior view, box defines area of image D. —D. Detail of posterioropisthosoma. —E. Anterior opisthosoma. —F. Spinnerets (arrow indicates absence of triad). Scale bars: A, C, E = 100 µm, B, D, F = 10 µm.



Pauly A) — cl013/f1 (Female), cl013/m1 (Male); AntsirananaProvince, P. N. Mt. D’Ambre, 12°32′S, 49°10′E, 1100, 23–28Nov 1993 (coll. Coddington et al.) — cl045/f1 (Female),cl045/m1 (Male); Tamatave (Toamasina), Ranotsara, 18°8′S,49°24′E, 10, forest reserve, littoral forest on sand, 18 October1993 (coll. Pauly A) — cl012/f1 (Female), cl012/j1 (Imma-ture), cl012/m1 (Male); Toamasina, Tamatave (Toamasina),18°8′S, 49°24′E, 10 (coll.) — cl022/f1 (Female); Antsiranana

Prov., 8.4 km NNW Manantenina, Marojezy Res., 14°26′S,49°45′E, 40–140, 10 November 1993 (coll. Coddington et al.)— cl046/f1 (Female), cl046/m1 (Male); Toamasina Prov.,50 km W Moramanga, 18°54.35′S, 47°53.37′E, 1300, 1 August1992 (coll. V Roth) — cl003/f1 (Female), cl004/f1 (Female).Etymology. Simon (1894: 757) reports the type specimensource as ‘Madagascar: Tamatave (Perrot)’. Presumably, henamed the species after its collector Perrot.

Fig. 18 A–F. Clitaetra episinoides, male left pedipalp, from Comoro Islands (cl6/m2): —A. Ectal. —B. Bulb, ectal. —C. Meso-apical. —D. Bulb,meso-apical. —E. Posterior. —F. Embolic conductor, ecto-ventral. — Scale bars = 20 µm.



Diagnosis. Clitaetra perroti females differ from all otherClitaetra species, except C. episinoides, by the presence of a‘tongue’ epigynum (Figs 19C–F and 22A,B). Clitaetra perrotiis approximately half to two thirds the size of C. episinoides.Further, the following combination of features separatesC. perroti females from C. episinoides: epigynum an opaquetongue, with spermathecae rarely visible through the cuticleanterior to the tongue (Fig. 19C–F); spermathecae separatedfrom the posterior tongue edge by one spermatheca diameteror less (Fig. 19E,F); venter medially white (Fig. 19B); cara-pace and opisthosoma broader than in C. episinoides (carapacelength to width ratio 1.1, opisthosoma length to width ratio1.1). A unique feature of C. perroti are the aggregate glandspigots of the female posterior lateral spinneret triad, whichare apart from the flagelliforms (Fig. 22F). Clitaetra perrotimales differ from all other Clitaetra species by the subtegu-

lum distally fused to the tegulum (Figs 20E and 24A,C).Clitaetra perroti males further differ from C. episinoides bynumerous denticles distally on embolus base (Fig. 24D,E), bythe broader sperm duct in the tegulum (Fig. 20E,G), andby the shorter embolic conductor (the ratio EC length tocymbium length less than 1.4 : 1).Description. Female (cl15, from Ambozotany; Figs 19 and21,22): Total length 5.7. Prosoma (Fig. 19A) 2.1 long, 1.7wide, 1.3 high; light brown. Sternum 0.95 long, 0.95 wide;medially light brown, laterally, caudally somewhat darker(Fig. 19B), with irregular patches of white pigment. AMEdiameter 0.11, ALE 0.14, PME 0.11, PLE 0.13. AME separ-ation 0.14, PME separation 0.19, PME–PLE separation0.18, AME–ALE separation 0.04, AME–PME separation0.21, ALE–PLE separation 0.20. Both eye rows recurved(Figs 19A and 21A,B). Clypeus height 0.15. Chelicerae

Fig. 19 A–F. Clitaetra perroti, femalemorphology, from Madagascar: A–E. Cl15.—A. Habitus, dorsal. —B. Habitus, ventral.—C. Epigynum, ventral. —D. Epigynum,posterior. —E. Epigynum, cleared, dorsal.—F. holotype (cl22), epigynum, ventral.Scale bars :A, B = 1.0 mm, C–F = 0.1 mm.



(Fig. 21E) with three prolateral, three retrolateral teeth, andapproximately 15 denticles. Cheliceral boss with approxi-mately 50 striae. Appendages. Legs yellow-light brown, withpatellae, tibiae, metatarsi, tarsi distally darker brown. Leg Ilength 10.1 (Fe 2.8, Pa 0.8, Ti 2.4, Me 2.7, Ta 1.3). Legformula 1-2-4-3. Opisthosoma (Fig. 19A,B) 3.9 long, 3.0 wide,2.4 high, oval, widest in middle, with three conspicuous pairsof dorsal apodemes, fourth inconspicuous. Dorsum whitish,with dark median band broken posteriorly (Fig. 19A), poste-rior part darker. Venter (Fig. 19B) with brown V-shapedmark, medially white, caudally with two pairs of white dots,pair of brown patches laterally to booklung covers. Sevenpairs of ventromedian apodemes, paired row of lateralsclerotizations. Spinnerets (Fig. 22D–F) typically nephiline(Hormiga et al. 1995), except the ectal aggregate gland spigotsof PLS triad apart from ‘flagelliform + mesal aggregate’pair (Fig. 22F). Epigynum as in Figs 19C–F and 22A,B. Epigy-

nal plate dark brown, opaque. Epigynum ventrally withtongue, copulatory openings on posterior sclerotization.Inner epigynum simple (Fig. 19E), with spermathecaeconnected to copulatory opening by tubular copulatoryduct.

Male (cl15, from Ambozotany; Figs 20 and 23,24): Totallength 2.9. Prosoma (Figs 20A–D and 23A,B) 1.3 long, 1.2wide, 0.8 high; light brown. Sternum 0.76 long, 0.69 wide;medially light brown, laterally, caudally somewhat darker(Fig. 20D). AME diameter 0.09, ALE 0.07, PME 0.07, PLE0.07. AME separation 0.08, PME separation 0.13, PME–PLE separation 0.11, AME–ALE separation 0.04, AME–PME separation 0.12, ALE–PLE separation 0.14. Both eyerows recurved (Figs 20A–C and 23A,B). Clypeus height 0.13.Chelicerae with three prolateral, three retrolateral teeth,approximately 10 denticles. Cheliceral boss with around 30striae. Appendages. Legs orange-brown, tibiae distally darker.

Fig. 20 A–G. Clitaetra perroti, malemorphology, from Madagascar (cl15/m1–2):—A. Habitus, dorsal. —B. Same, ectal.—C. Same, frontal. —D. Same, ventral. —E.Left pedipalp, ectal. —F. Same, mesal. —G.Left bulb, transparent, showing spermduct, ectal. Scale bars: A–D = 1.0 mm, E–G = 0.1 mm.



Leg I length 8.0 (Fe 2.3, Pa 0.6, Ti 2.0, Me 2.3, Ta 1.0). Legformula 1-2-4-3. Opisthosoma (Figs 20A,B,D and 23C–F) 1.7long, 1.2 wide, 0.9 high, oval, truncated posteriorly, withscutum, three pairs of dorsal apodemes. Dorsum whitish, withdark median band, broken posteriorly (Fig. 20A). Venterbrown to dark brown, caudally with two pairs of white dotsaround spinnerets (Fig. 20D). Ventral apodemes, sclerotiza-

tions as in female (Fig. 23C). Pedipalp as in Figs 20E–G and24A–F: globular tegulum, subtegulum distally fused to tegu-lum (Figs 20E and 24A,C), long, narrow embolic con-ductor enveloping embolus. Embolus base distally withnumerous denticles (Figs 20F and 24D–F).Variation. Females: prosoma length ranges from 1.9 to 2.6(n = 14); total length from 4.4 to 7.0 (n = 14). Prosoma and

Fig. 21 A–F. Clitaetra perroti, female, from Madagascar (cl15/f2): —A. Prosoma, dorsoectal. —B. Head region, dorsofrontal. —C. Prosoma, ventral,box delimits area of image D. —D. Detail of sternum with slit sensilla (arrow) adjacent to coxa 3. —E. Chelicerae and maxillae, ventroectal.—F. Tarsus 4 showing claws and sustentaculum (partly hidden); note long median claw (arrow). Scale bars: A–C, E = 100 µm, D, F = 10 µm.



legs vary in coloration from dull grey or light yellow tored-brown. In some females the carapace exhibits threealmost parallel darker bands. Irregular patches of whitepigment are present on sternum of most preserved females.Some females have the fourth (smaller) tooth on cheliceralretromargin but most have only three. Cheliceral denticlenumbers 13–26. Cheliceral boss striae from 50 to more than100. Dorsal pattern varies in contrast and coloration, but is

generally as illustrated (Fig. 19A). Venter pattern is variablebut always has the conspicuous white median part, with theexception of the females from Mt. D’Ambre in northernMadagascar, which have a dull grey venter. Some females(including the holotype) lack the dark V mark around thewhite median part, but instead have a darker posterior venterthan illustrated. At least five, but mostly seven pairs ofventromedian apodemes are present, and a paired row of

Fig. 22 A–F. Clitaetra perroti, female opisthosoma, from Madagascar (A–C. Cl13/f1. D–F. cl15/f2): —A. Epigastric area, ventral. —B. Same,posterior. —C. Anterior dorsum detail. —D. ALS. —E. PMS. —F. PLS. Scale bars: A, B = 100 µm, C–E = 10 µm.



ventrolateral sclerotizations. Epigynal tongue can be blackand opaque (Fig. 19C) as in females from Ambozotany, orlighter brown with spermathecae visible through the cuticle,as in females from other localities including the holotype(Fig. 19F).

Males: prosoma length ranges from 1.3 to 1.7 (n = 4), totallength from 2.9 to 3.4 (n = 4). Second pair of dorsal apodemes

is not apparent in all males. In some males the abdominalscutum is not pronounced. Specimens vary in coloration fromdull grey or light yellow to orange.Distribution. Madagascar (Fig. 26).Natural history. The label data suggest that the speciesinhabits sand forests, rain forests (Ranomafana), and intro-duced pine forests of Madagascar.

Fig. 23 A–F. Clitaetra perroti, male somatic morphology, from Madagascar (cl13/m1): —A. Prosomal head region, dorsal. —B. Prosoma, lateral (notemesal orientation of palpal cymbium). —C. Opisthosoma, ventral, box defines area of image D. —D. Epigastric region with four epiandrous glandspigots (arrows). —E. Lateral opisthosoma detail. —F. Frontal opisthosoma detail (with sensory setae). Scale bars: A–C = 100 µm, D–F = 10 µm.



Clade Indoetra subgen. nov.

Etymology. A combination of Indo- (for Indian subcontinent)and Clitaetra; the name is feminine in gender.Monophyly. The clade is currently represented by a singlespecies; see Discussion.Phylogenetic definition. The subgenus Indoetra is defined as

the least inclusive clade with C. thisbe Simon, 1903 and anyother discovered Asian Clitaetra species. See Discussion.Diagnosis. Epigynum a transverse plate with a transparentarea anteriorly (Fig. 25B,C). Small spiders (3.5 mm).Composition. The clade currently contains a single species,C. thisbe.Type species. Clitaetra thisbe Simon.

Fig. 24 A–F. Clitaetra perroti, male left pedipalp, from Madagascar (cl3/m1): —A. Ectal. —B. Paracymbium detail showing margin fold (arrow).—C. Posterior-ectal. —D. Mesal, showing embolus base denticles (arrow). —E. Apical, showing embolus base denticles (arrow). —F. Ventral.Scale bars = 20 µm.



Distribution. Sri Lanka (Fig. 26).

Clitaetra thisbe Simon, 1903 (Fig. 25)

Clytaetra thisbe Simon, 1903: 24, description of female (fromSri Lanka). Female holotype in MNHN, labelled ‘16296Clit. thysbe E.S. Galle’, examined. The type locality (fromSimon 1903) is Galle in Sri Lanka (approximately 6.02′N,80.13′E).Clitaetra thisbe: Simon 1907: 284.Clytaetra thisbe: Roewer 1942: 925.Clitaetra thisbe: Bonnet 1956: 1100; Platnick 2005.Additional material examined. None.Etymology. Unknown.Diagnosis. At 3.5 total female length, this species is by far thesmallest in the genus. The female epigynum (Fig. 25B,C)differs from all other species: a simple transverse sclerotizededge with copulatory openings posteriorly, and with thecopulatory ducts, fertilization ducts, and spermathecae clearlyvisible through the ventral cuticle.Description. Female (holotype; Fig. 25): Total length 3.5.Prosoma 1.6 long, 1.3 wide, 0.9 high; yellow-brown. Sternum0.7 long, 0.6 wide; yellow, with two pairs of sternal slit sensilla.AME diameter 0.13, ALE 0.13, PME 0.08, PLE 0.12. AMEseparation 0.09, PME separation 0.14, PME–PLE separation0.17, AME–ALE separation 0.04, AME–PME separation0.15, ALE–PLE separation 0.15. Both eye rows recurved.Clypeus height 0.03. Chelicerae with three prolateral, threeretrolateral teeth, approximately 15 denticles. Cheliceralboss with 40–50 striae. Appendages. Legs yellow-brown. LegI length 7.8 (Fe 2.1, Pa 0.6, Ti 1.7, Me 2.1, Ta 0.9). Leg formula1, 2 = 4, 3. Opisthosoma 2.0 long, 1.7 wide, 1.1 high. Opisthosoma

oval, dorsum yellow-brown with grey pattern, four pairs ofmedian apodemes (Fig. 25A). Venter grey, median part lighter,with five pairs of ventromedian apodemes. Epigynum a simpletransverse sclerotization (Fig. 25B,C). Inner epigynum simple,as diagnosed.

Male: Unknown.Distribution. Known only from the type locality in Sri Lanka.This is the only species of the genus so far known to occuroutside of Africa and Madagascar including its adjacentislands (Fig. 26).Natural history. Unknown.

Fig. 25 A–C. Clitaetra thisbe, female holo-type from Sri Lanka: —A. Habitus, dorsal.—B. Epigynum, ventral. —C. Epigynum,posterior. Scale bars: A = 1.0 mm, B, C = 0.1 mm.

Fig. 26 Clitaetra species distribution map. Note that C. simoni andC. thisbe are only known from the type localities, D. R. Congo andSri Lanka, respectively.



DiscussionClassificationThis study corroborates the monophyly of Clitaetra, Heren-nia, Nephila and Nephilengys, all classically considerednephilines. The nephiline placement as sister to other arane-oids minus araneids disputes all previous hypotheses. Suchplacement will be further tested with a denser taxonomicsample. However, it now seems clear that Nephilinae asdefined by Hormiga et al. (1995) is polyphyletic, does notcontain Phonognatha, and does not group with tetragnathids.Therefore, I propose a revised classification elevating theclade (Clitaetra + (Herennia + (Nephila + Nephilengys))) tofamily rank, Nephilidae. The newly defined subfamilyNephilinae is proposed to contain the clade (Herennia +(Nephila + Nephilengys)). The remaining nephilid clade con-taining all Clitaetra species is here proposed to be treatedas another subfamily, Clitaetrinae, which is currently ident-ical in composition and phylogenetic definition to the singlegenus within (Clitaetra), a so called Gregg’s Paradox (e.g. Ruse1971).

In a rankless classification system such as the PhyloCode(Cantino & de Queiroz 2004) Gregg’s Paradox does not exist.There has been a vigorous debate on the pros and cons ofrankless vs. Classical Linnaean classification systems (areview is in preparation). In a rankless system, a new cladename does not have to be accompanied by naming theremaining clades, but in classical Linnaean classification itdoes. For example, the only well supported clitaetrine clade(Fig. 27B) contains the three currently known (and probablyseveral unknown) mainland African species (C. irenae +(C. clathrata + C. simoni)). It makes some sense to name the clade,simply for the sake of discussing the interesting biologicalaspects it offers (biology, biogeography, anatomy, etc.). How-ever, naming a species group in the Linnaean classificationsystem requires a rank, and if the new name, Afroetra, receivesthe genus rank, one would have to also name, at the genusrank, the two clades with two (C. episinoides + C. perroti ) anda single species (C. thisbe), respectively, to avoid their para-phyly. This study uses Linnaean ranks, but to some extentavoids disruptive changes in species nomenclature by treatingthe three newly named clades (Afroetra, Clitaetra, Indoetra)arbitrarily at the subgenus level. To follow rankless classifica-tion (Cantino & de Quieroz 2004), the clade Afroetra couldbe newly named and defined, with no forced adjacent newclade names and no disruption to Clitaetra species nomenclature.Thus, another Gregg’s Paradox, this time in the monotypicsubgenus Indoetra, would be avoided. A simple alternative inthe Linnaean system, of course, is not to name new clades,and if they do merit discussion, give them informal names,a standard practice in recent araneological phylogeneticliterature (Scharff & Coddington 1997; Griswold et al.1998; Agnarsson 2004, 2005; Agnarsson & Kuntner 2005).

However, none of the alternatives provides a formal lineagename. The biogeographic implications of basal nephilids domerit discussion and more research and thus the (subgeneric)names Afroetra, Indoetra and Clitaetra are used here.

Fig. 27 A, B. Phylogenetic results: —A. Strict consensus of eightshortest cladograms (L = 531, CI = 42, RI = 70) collapsing threenodes. —B. Successively weighted tree identical to one of thefundamental cladograms, representing a working hypothesis ofnephilid relationships. Clade support given as bootstrap (abovebranch) and Bremer index (below branch).



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BiogeographyNephilid spiders recorded from the Dominican amberinclude Nephila (Wunderlich 1986) and a number of unplacedfossil genera (Wunderlich 2004), but no clitaetrine fossilsare known, and no clitaetrines (or Herennia) have been re-corded from the neotropics. Clitaetra cladistic biogeography(Fig. 29) suggests the area cladogram ((Africa + Madagascarwith adjacent islands) + Indian subcontinent), which does notagree with the currently understood continental drift theory(Brown & Lomolino 1998; McLoughlin 2001), becauseIndian and Madagascar species do not group. If the Gondwa-nan distribution of Clitaetrinae is explained solely by vicari-ance with no dispersal events, the phylogeny allows roughclade age estimations (in Fig. 29 minimum clade age is given).The west–east Gondwana split separating S America + Africafrom the remainder of Gondwana (including Madagascar and

India) is estimated at 160 Myr. The monophyly of the Africanand Madagascar species would suggest that the clade predatesthe split. Thus, assuming no intercontinental dispersalevents, the clade Afroetra + Clitaetra must be at least 160 Myrold, and its sister clade Indoetra, by definition, of the sameage. All more inclusive clades can be assumed to be older.Thus, the basal clitaetrine split between the Indian andMadagascar (plus African) species probably well predates thesplit between India and Madagascar (60 Mya), and its sisterclade, Nephilinae, must be of the same age (more than 160Myr old). Likewise, the nephilid clade (and its sister cladewith the remaining araneoids) cannot be less than 160 Myrold, predating the separation of (Madagascar + India) from(Australia + Antarctica) (125–130 Mya).

Given the above estimates of clade ages it should come asno surprise that the basal nephilid dichotomy and the deep

Fig. 29 Nephilid relationships with Clitaetrageographic distributions, simplified after thesuccessively weighted phylogeny. Proposedclassification combines Linnaean ranks withphylogenetic clade definitions. Minimum ageof the clade Afroetra + Clitaetra estimated bythe split of west and east Gondwana (see text).



araneoid relationships (see also Kuntner 2005) are poorlysupported due to lack of solid morphological and behaviouralsynapomorphies.

ConclusionsThe biology of Clitaetra remains insufficiently known. Theclade Clitaetrinae is sister to all other nephilid lineages(Figs 27–29) and thus its scored features will affect the nephi-lid character polarizations. A potential rediscovery of Clitae-tra thisbe, not recorded since the original description from SriLanka, would enable testing as to whether the morphologicaland behavioural data here presented hold also for the basal-most clitaetrine species. Unfortunately, the type locality ofC. thisbe (Galle, Sri Lanka) has been heavily deforested in thelast century and was lately disastrously affected by the 2004tsunami. However, it is likely that there are further (Indoetra)species on the Indian subcontinent awaiting discovery.

AcknowledgementsI thank J. Coddington, G. Hormiga, , I. Agnarsson andJ. Miller for their help, encouragement and comments. M.Allard, J. Clark, D. Lipscomb and C. Thompson made com-ments on an early draft, which formed a chapter of my PhDthesis. I. Agnarsson patiently read and discussed several draftsof this paper, and further useful comments were provided bytwo reviewers. F. Alvarez-Padilla, L. Lopardo, D. De Rocheand S. Larcher offered assistance and help; S. Whittaker andP. Herendeen provided SEM help, and K. Darrow kindlyhelped with digital image manipulation. I thank the followingcurators and collection managers for their kind assistancewith loans and/or curation: C. Griswold and D. Ubick atCAS, G. Doria at MCSNG, G. Giribet and L. Leibenspergerat MCZ, C. Rollard at MNHN, A. Dippenaar-Schoeman atPPRI, R. Jocqué at RMCA, J. A. Coddington, S. Larcher andD. M. De Roche at USNM. A. and J. Leroy kindly sharedtheir useful unpublished data on Clitaetra irenae naturalhistory and distribution, and R. Jocqué provided his un-published photograph of Clitaetra episinoides. I thank J.Coddington for patiently taking over from me in the Africanfield at night to record spider behaviour when fatigue kickedin. The fieldwork in South Africa, conducted jointly withI. Agnarsson, M. Arnedo, J. Coddington, G. Hormiga andM. Ramirez, was made possible by the help of M. Hamer,T. Crouch, C. Whitmore, L. Prendini, A. Dippenaar-Schoeman,N. Larsen and A. Kilpin. The staff at Phinda ResourceReserve are kindly acknowledged for their hospitality. Thisproject was supported by the U.S. National Science Founda-tion (PEET grant DEB-9712353 to Hormiga and Codding-ton), by the Sallee Charitable Trust (grant to Agnarsson andKuntner) and the OTS-STRI-Mellon Research ExplorationAward (to Kuntner and Sereg). I further acknowledge mate-rial and financial support of George Washington University,

Smithsonian Institution, the Ministry of Science of theRepublic of Slovenia and the Biological Institute of the Slove-nian Academy of Sciences and Arts. Finally, this long projectwould have been impossible without the patience of my wifeIrena, to whom I dedicate this work.

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Preston-Mafham, R. (1991). The Book of Spiders. Edison: ChartwellBooks.

Robinson, M. H. & Lubin, Y. D. (1979). Specialists and generalists:the ecology and behavior of some web-building spiders fromPapua New Guinea, 1. Herennia ornatissima, Argiope ocyaloides andArachnura melanura (Araneae: Araneidae). Pacific Insects, 21, 97–132.

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Scharff, N. & Coddington, J. A. (1997). A phylogenetic analysis ofthe orb-weaving spider family Araneidae (Arachnida, Araneae).Zoological Journal of the Linnean Society, 120, 355–434.

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Wunderlich, J. (1986). Spinnenfauna gestern und heute. Fossile Spinnenin Bernstein und Ihre heute lebenden Verwandten. Wiesbaden: EricBauer; Quelle & Meyer.

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Appendix 1. List of characters and character statesList of characters and character states used in the phyloge-netic analysis, described further in the treatments of Herennia(Kuntner 2005) and Nephilengys (Kuntner unpublished). Arigorous homology test of nephilid characters will be pre-sented elsewhere (Kuntner, Hormiga and Coddington,unpublished).1. Female cephalic region: (0) low (Fig. 9A); (1)conspicuously high.2. Female carapace: (0) piriform (Fig. 12B); (1) oval withwide head region.3. Female carapace edge: (0) smooth; (1) ridged (Kuntner2005: fig. 5D).4. Female carapace edge: (0) glabrous or with few hairs;(1) with an extensive row of hairs (Fig. 9A,B).5. Female median pair of prosomal tubercles: (0) absent;(1) present.6. Fovea (female carapace): (0) inconspicuous; (1)pronounced.7. Female carapace macrospines: (0) absent; (1) present.8. Female carapace warts: (0) absent; (1) present (Kuntner2005: fig. 17).9. Female carapace hairs: (0) present (Fig. 1A); (1) absent.10. Female median eye region: (0) rounded; (1) medianeyes on a tubercle.11. Female lateral eye region: (0) lateral eyes on separ-ate tubercles; (1) lateral eyes on a single tubercle; (2)rounded.12. Female LE separation from ME: (0) not widelyseparated; (1) widely separated.13. Posterior eye row (dorsal view): (0) straight torecurved; (1) procurved.14. Female PME: (0) less than one PME diameter apart; (1)one PME diameter or more apart.15. Female PLE size: (0) equal or less than PME; (1) largerthan PME.16. PME canoe tapetum: (0) absent; (1) full; (2) narrow.17. PLE canoe tapetum: (0) absent; (1) full; (2) narrow.18. Female clypeus height: (0) low (less than three AMEdiameters); (1) equal or more than three AME diameters.19. Endites: (0) very long (> 2× width); (1) short (length < 2×width).20. Labium and sternum: (0) separate; (1) fused.21. Female sternum: (0) longer than wide; (1) as wide orwider than long.22. Sternal slit sensilla: (0) present (Fig. 3F); (1) absent.23. Female sternum colour pattern: (0) inconspicuouslycoloured; (1) uniformly orange/red; (2) medially dark,laterally pale; (3) medially light, laterally dark; (4) with yellowspots corresponding to tubercles.24. Sternal white pigment: (0) absent; (1) present(Figs 12C and 19B).

25. Female sternal tubercle I: (0) absent; (1) present(Kuntner 2005: fig. 6E).26. Female sternal tubercle II: (0) absent; (1) present.27. Female sternal tubercle III: (0) absent; (1) present.28. Female sternal tubercle IV: (0) absent; (1) present.29. Female frontal sternal tubercle: (0) absent; (1) present.30. Frontal sternal tubercle: (0) small; (1) large projection.31. Female chilum: (0) absent; (1) present.32. Female chelicerae: (0) massive (width > 1/2 length); (1)slender (width < 1/2 length).33. Cheliceral ectal margins: (0) smooth; (1) withstridulatory striae.34. Cheliceral boss: (0) present (Fig. 9C,D); (1) absent.35. Cheliceral boss surface: (0) smooth; (1) striated(Figs 3E and 9C,D).36. Prosomal supracheliceral lobe (PSL): (0) present(Fig. 3E); (1) absent.37. Cheliceral furrow: (0) denticulated (Fig. 3D); (1) smooth.38. Female first femur: (0) more/less straight; (1) sigmoidal.39. Femoral macrosetae: (0) present; (1) absent.40. Femoral (I, II) macrosetae length: (0) long; (1) short,stout.41. Female femur I group of prolateral long spines: (0)absent; (1) present (Kuntner 2005: fig. 31B).42. Spine socket colour: (0) same as legs; (1) dark (Fig. 12A).43. Dorsal femoral trichobothria: (0) absent; (1) present.44. Female tibia I tufts: (0) absent; (1) present.45. Female tibia II tufts: (0) absent; (1) present.46. Female tibia IV tufts: (0) absent; (1) present.47. Patella-tibia autospasy: (0) absent; (1) present.48. Ventral tarsus IV setae: (0) irregular; (1) comb-like.49. Tarsus 4 median claw: (0) long (as long or longer thanthe main claw; Fig. 21F); (1) short (shorter than the pairedmain claw).50. Sustentaculum: (0) present (Fig. 14F); (1) absent.51. Sustentaculum angle: (0) wide, diverging from othersetae; (1) narrow, parallel to other setae (Fig. 14F).52. Female abdomen length: (0) very long (> 2× width); (1)long (longer than wide, but < 2× width); (2) Short (as wide aslong or wider).53. Female abdomen width: (0) elliptical; (1) widestanteriorly; (2) widest posteriorly; (3) pentagonal (Fig. 7A).54. Female lateral abdominal margin: (0) smooth; (1) with3–4 pairs of lobes.55. Female anterior abdominal humps: (0) absent; (1)present.56. Female abdomen tip: (0) rounded; (1) truncated(Fig. 10A).57. Female ventromedian sclerotizations: (0) absent; (1)paired sclerotizations (Figs 4A and 10A).58. Female ventromedian sclerotizations: (0) 1–5 pairs;(1) 6–11 pairs (Figs 4A and 10A).



59. Female ventrolateral abdominal sclerotizations: (0)present (Figs 4A,B and 10A,B); (1) absent.60. Ventrolateral abdominal sclerotizations: (0) onepaired line of small dots; (1) sclerotizations in severallines.61. Female dorsomedian abdominal apodemes: (0)absent; (1) 3–5 prominent pairs.62. Female dorsolateral abdominal sclerotizations: (0)present; (1) absent.63. Female dorsocentral abdominal sclerotizations: (0)absent; (1) present.64. Female abdominal sigillae: (0) absent; (1) present.65. Female anterior abdomen: (0) without; (1) with abroad light-pigmented band.66. Female abdominal dorsal pattern: (0) inconspicuous;(1) conspicuous.67. Female dorsum dark spots: (0) absent; (1) present.68. Female dorsum paired light spots: (0) absent; (1)present.69. Female dorsum ‘butterfly’ pattern: (0) absent; (1)present (Figs 1A and 6A).70. Female dorsum ‘grid’ pattern: (0) absent; (1) present(Figs 1A and 6A).71. Female abdomen tip colour: (0) no different to thesubapical abdomen; (1) paired white dots around spinnerets(Fig. 12C).72. Female abdomen silver pigment spots: (0) absent; (1)present.73. Female venter light pigmented pattern: (0) absent; (1)present.74. Female venter light pigmented pattern form: (0) onecentral light area (Fig. 19B); (1) transverse line(s); (2) fourlarge spots; (3) numerous spots; (4) longitudinal lines.75. Booklung cover: (0) grooved; (1) smooth.76. Area around female booklung spiracle: (0) littlesclerotized; (1) strongly sclerotized.77. Posterior epigynal plate: (0) round; (1) grooved.78. Epigynal ventral area: (0) low; (1) swollen.79. Epigynal posterior area: (0) round; (1) tongue-shaped(Figs 12E,F and 15B).80. Epigynal openings: (0) simple (Fig. 7C); (1) in chambers.81. Chamber opening position: (0) medial; (1) lateral.82. Epigynal septum: (0) absent; (1) present.83. Epigynal septum shape: (0) simple border betweenchambers; (1) extensive, broader posteriorly; (2) extensive,broader anteriorly.84. Epigynal paired sclerotized pocket: (0) absent; (1)present.85. Anterior epigynal area: (0) with a pair of apodemes; (1)round.86. Cuticle anterior to the epigynal area: (0) rounded; (1)depressed.

87. Epigynal anterior area: (0) round; (1) depressed(Fig. 7B, EAD).88. Epigynal lateral area: (0) round; (1) curved (Fig. 7B,ELC).89. Copulatory opening position: (0) caudal (Fig. 7B,C);(1) ventral.90. Caudal copulatory openings: (0) on the posteriorsclerotized epigynal margin (Fig. 1C,E); (1) anterior to theposterior margin (Fig. 7C).91. Copulatory opening form: (0) elongated slit openings;(1) rounded openings (Fig. 7C).92. Copulatory duct morphology: (0) flattened duct (longerthan wide, flat); (1) tube (longer than wide, cylindrical); (2)broad attachment to body wall (wider than long).93. Spermathecae: (0) lobed; (1) spherical; (2) oval.94. Spermathecae separation: (0) wide (separated morethan 2 widths); (1) small or none (separated less than twowidths; Fig. 7D).95. Epigynal sclerotized arch: (0) absent; (1) present.96. Female copulatory aperture: (0) never plugged; (1)sometimes plugged with emboli a/o conductors.97. Female copulatory plugs: (0) emboli; (1) emboli plus(embolic) conductors98. Cribellum: (0) present; (1) absent.99. ALS piriform gland spigot bases: (0) normal (Fig. 4E);(1) reduced.100. PMS nubbin: (0) absent; (1) present (Fig. 4F).101. PMS aciniform field: (0) extensive; (1) sparse (Fig. 4F).102. PLS mesal cylindrical gland spigot base: (0)subequal to other PLS cylindrical spigot (Fig. 4F); (1) larger.103. PLS mesal cylindrical gland spigot position: (0)central; (1) peripheral (Fig. 4F).104. PLS aggregate-flagelliform relation: (0) aggregatesapart from flagelliform (Fig. 25D–F); (1) distal aggregatespigots embrace flagelliform (Figs 4F, 10F and 15F).105. PLS aggregate gland spigot: (0) normal (Fig. 15F);(1) large.106. Male size: (0) more than half the size of female; (1) lessthan 0.4 female.107. Male dorsal abdomen: (0) cuticle soft; (1) with scutum(Fig. 17C–E).108. Male lateral eyes: (0) separate (Fig. 16A,E); (1)juxtaposed.109. Male cephalic region: (0) narrower than in female; (1)same proportion to cephalothorax as in female.110. Male clypeus: (0) as in female; (1) more horizontal.111. Male v. female cheliceral size: (0) same; (1) larger; (2)smaller.112. Male paturon posteriorly: (0) smooth; (1) with atubercle.113. Male leg II tibial macrosetae: (0) similar to those ontibia I; (1) stronger and more robust; (2) absent.



114. Male endite depression: (0) absent; (1) present.115. Male palpal trochanter: (0) short (twice the width orless); (1) long (more than twice the width).116. Male palpal femoral tubercle: (0) absent; (1) present.117. Male palpal patella macrosetae: (0) none; (1) one; (2) two.118. Male palpal tibia length: (0) short (not exceeding 1.5×width); (1) long (exceeding 1.5× width).119. Cymbium length: (0) short (less than 2× width); (1)long (more than 2× width).120. Cymbium orientation: (0) dorsal; (1) mesal (Fig. 23B).121. Cymbial ectal margin: (0) sclerotized as cymbium; (1)transparent.122. Paracymbium (P): (0) absent; (1) present (Figs 2A–C,5D,F, 8C, 13A,D and 18A,B).123. Paracymbial base sclerotization: (0) like cymbium;(1) less sclerotized.124. Paracymbium morphology: (0) short basal structure,more or less hook-shaped; (1) longer than wide and finger-like; (2) flat and roughly rectangular; (3) U-shaped; (4) flatand roughly triangular; (5) Phonognatha condition.125. Paracymbium edge: (0) glabrous; (1) with setae.126. Anterior paracymbial apophysis (APA): (0) absent;(1) present.127. Paracymbial margin fold: (0) absent; (1) present.128. Paracymbium apically: (0) rounded; (1) with a prong.129. Tegulum in ectal view: (0) same size as or larger thansubtegulum; (1) smaller than subtegulum.130. Reservoir course: (0) spiralled; (1) with a switchback(Fig. 2B,C).131. Ventral tegular switchback: (0) single (Fig. 2C); (1)double (Fig. 2D).132. Ejaculatory duct: (0) within the entire length ofembolus; (1) joins distal embolus.133. Median apophysis (MA): (0) absent; (1) present.134. Median apophysis: (0) without sperm duct; (1) with aloop of the sperm duct.135. Median apophysis thread-like spur: (0) absent; (1)present.136. Apical tegular apophysis (ATA): (0) absent; (1) present.137. Ventral tegular apophysis (VTA): (0) absent; (1)present (Fig. 2).138. Mesal tegular apophysis (MTA): (0) absent; (1) present.139. Theridiid tegular apophysis (TTA): (0) absent; (1) present.140. Conductor (C): (0) present; (1) absent.141. Conductor size: (0) small (less than half bulb volume);(1) large (more than half bulb volume).142. Conductor form: (0) rounded; (1) grooved for embolus.143. Conductor and embolus association: (0) separate; (1)embolus enclosed in conductor; (2) embolus and conductorspiral.144. Embolic conductor (EC): (0) absent; (1) present(Figs 2, 5D,E, 8,11C,E,F, 13, 18, 20E–G and 24A,C–F).

145. EC membrane: (0) absent; (1) present.146. EC shape: (0) complex; (1) finger-like.147. Finger EC: (0) short; (1) long.148. EC (division): (0) not subdivided; (1) subdivided intomore sclerites.149. EC edge: (0) smooth; (1) ridged.150. EC curvature: (0) more/less straight; (1) sigmoidal; (2)bent distally.151. EC tip: (0) straight; (1) with a hook.152. EC subdistal protuberance: (0) absent; (1) present.153. Embolus (E) length: (0) long (> 2× CB); (1) medium(0.5–1.5 CB length); (2) Short (< 1/2 cymbium).154. Embolus form: (0) thin (Fig. 13C); (1) thick; (2) filiform.155. Embolus-tegulum orientation: (0) parallel; (1) 90 deg.156. Embolus-tegulum membrane: (0) absent; (1) present.157. Embolus base: (0) thin; (1) enlarged (= radix).158. Embolus base distal part: (0) smooth; (1) denticulated(Fig. 24D,E, arrow).159. Embolic apophysis: (0) absent; (1) present.160. Radical membrane: (0) absent; (1) present.161. Stipes: (0) absent; (1) present.162. Embolus constriction: (0) absent; (1) present.163. Embolus: (0) smooth; (1) hooked.164. E distal apophysis: (0) present; (1) absent.165. Embolus tip: (0) flat; (1) cylindrical.166. Web architecture: (0) orb; (1) sheet; (2) gum foot.167. Orb-web angle: (0) horizontal (0–45 deg); (1) vertical(46–90 deg).168. Orb shape: (0) round; (1) rectangular (Fig. 10).169. Silk colour: (0) white; (1) golden.170. Stabilimentum: (0) absent; (1) present.171. Barrier (3D) web: (0) absent; (1) present.172. Hub position: (0) aerial; (1) against substrate.173. Hub relative position: (0) central; (1) displaced up; (2)displaced down.174. Hub bite-out: (0) present; (1) absent.175. Hub: (0) closed; (1) open.176. Hub-cup: (0) absent; (1) present.177. Hub loop — nonsticky spiral transition: (0) gradual;(1) abrupt.178. Radius construction: (0) cut and reeled; (1) doubled.179. Radius attachment on frame: (0) attached singly; (1)attached twice.180. Secondary (split) radii: (0) absent; (1) present.181. Tertiary (split) radii: (0) absent; (1) present.182. Sticky spiral: (0) spiralling; (1) parallel.183. Non-sticky spiral: (0) removed; (1) persists in web.184. NSS form: (0) linear; (1) zig-zag Nephila form.185. First SS spiral-NSS contact: (0) NSS contacted; (1)no contact.186. Sticky spiral localization: (0) oL1; (1) iL1; (2) oL4.187. Web posture: (0) flexed legs I, II; (1) extended legs I, II.



188. Argiope posture: (0) absent; (1) present.189. Attack behaviour: (0) wrap-bite; (1) bite-wrap.190. Wrap-bite silk: (0) dry; (1) sticky.191. 191. Cheliceral clasp: (0) absent; (1) present.192. Bulbus detachment (eunuchs): (0) absent; (1)present.

193. Body shake: (0) absent; (1) present.194. Side change: (0) absent; (1) when in danger, rushing onother side of orb.195. Partial web renewal: (0) absent; (1) present.196. Retreat: (0) absent; (1) off-web; (2) in web.197. Retreat form: (0) silken tube; (1) utilization of a leaf.

Appendix 2. Phylogenetic matrixPhylogenetic matrix, available electronically from the author in Winclada and NONA (ss) formats. Unknown entries = ?,inapplicable data = –.

Characters

Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39

1 Deinopis 0 0 0 0 0 0 0 0 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 – 0 0 0 0 0 0 0 0 02 Uloborus 0 0 0 1 0 0 0 0 0 0 2 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 – 0 0 0 0 0 0 0 0 03 Araneus 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 1 0 1 0 0 0 0 0 1 1 1 0 1 0 1 0 0 0 0 0 0 0 04 Argiope aurantia 0 0 0 1 0 0 0 0 0 1 1 0 1 1 1 2 2 0 1 0 0 0 3 1 1 1 1 0 0 – 1 0 0 0 0 0 0 0 05 Argiope argentata 0 0 0 1 0 0 0 0 0 1 1 0 1 1 1 2 2 0 1 0 1 0 3 1 1 1 1 0 0 – 1 0 0 0 0 0 0 0 06 Linyphia 0 0 0 0 0 1 0 0 1 0 2 0 0 1 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 – 0 0 1 0 – 1 1 0 07 Pimoa 0 0 0 0 0 1 0 0 1 0 2 0 0 0 0 1 1 1 1 0 0 1 0 0 0 0 0 0 0 – 0 0 1 1 – 1 1 0 08 Steatoda 0 0 0 0 0 0 0 0 1 0 2 0 0 1 0 1 1 1 1 0 0 1 0 0 0 0 0 0 0 – 0 1 0 1 – 1 1 0 19 Nesticus 0 0 0 0 0 1 0 0 1 0 2 0 1 1 0 1 1 1 1 1 0 1 0 0 0 0 0 0 0 – 0 1 0 1 – 1 0 0 110 Epeirotypus 1 0 0 0 0 0 0 0 1 0 2 0 1 1 0 1 1 0 1 0 0 0 0 0 0 0 0 0 0 – 0 0 0 1 – 1 0 0 011 Tetragnatha 0 0 0 0 0 1 0 0 1 0 2 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 – 0 1 0 0 0 0 1 0 012 Meta 0 0 0 0 0 1 0 0 1 0 2 0 0 0 0 1 1 0 1 1 0 0 0 0 1 1 1 0 1 0 0 1 0 0 0 0 1 0 013 Leucauge 0 0 0 0 0 1 0 0 1 0 2 0 0 1 0 1 1 0 1 1 1 0 0 0 1 0 1 0 0 – 0 0 0 0 0 0 1 0 014 Deliochus 0 0 0 0 0 0 0 0 1 0 2 1 0 1 0 1 1 0 1 0 0 0 0 0 1 0 0 0 0 – 1 0 0 0 0 0 0 1 015 Phonognatha graeffei 1 1 0 0 0 0 0 0 1 1 2 1 0 1 0 1 1 0 1 0 0 0 3 0 1 0 1 0 0 – 1 0 0 0 0 0 0 1 016 Clitaetra episinoides 0 0 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 1 0 1 0 3 1 0 0 0 1 0 – 0 0 0 0 1 0 0 0 017 Clitaetra perroti 0 0 0 0 0 0 0 0 0 0 2 0 0 1 1 0 0 0 1 0 1 0 0 1 0 0 0 0 0 – 0 0 0 0 1 0 0 0 018 Clitaetra clathrata 0 0 0 1 0 0 0 0 0 0 2 0 0 1 1 0 0 0 1 0 1 0 3 1 0 0 0 0 0 – 1 0 0 0 1 0 0 0 019 Clitaetra irenae 0 0 0 1 0 0 0 0 0 0 2 0 0 1 1 0 0 0 1 0 0 0 3 1 0 0 0 0 0 – 0 0 0 0 1 0 0 0 020 Clitaetra simoni 0 0 0 1 0 0 0 0 ? 0 2 0 0 1 1 0 0 0 1 0 1 0 3 1 1 0 0 0 0 – 1 0 0 0 1 0 0 0 021 Clitaetra thisbe 0 0 0 0 0 0 0 0 ? 0 2 0 0 1 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 – 0 0 0 0 1 0 0 0 022 Nephila clavipes 1 1 ? 0 ? 0 0 0 0 1 1 1 0 1 1 1 1 0 1 0 0 0 2 0 1 1 1 0 1 0 1 0 0 0 1 0 0 0 023 Nephila fenestrata 1 1 1 1 1 0 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0 2 0 1 1 1 1 1 0 1 0 0 0 1 0 0 0 024 Nephila pilipes 1 1 0 0 ? 0 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0 0 0 1 1 1 0 1 0 1 0 0 0 1 0 0 0 025 Nephila antipodiana 1 1 1 1 1 0 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0 ? 0 1 1 1 1 1 1 1 0 0 0 1 0 0 0 026 Nephila clavata 1 1 0 1 0 0 0 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0 3 1 1 1 1 1 1 0 1 0 0 0 1 0 0 0 027 Nephila plumipes 1 1 ? 1 ? 0 0 0 0 1 1 1 0 1 ? 1 1 0 1 0 0 0 4 0 1 1 1 1 1 1 1 0 0 0 1 0 0 0 028 Nephila edulis 1 1 0 1 1 0 0 0 0 1 1 1 0 1 ? 1 1 0 1 0 0 0 4 0 1 1 1 1 1 0 1 0 0 0 1 0 0 0 029 Herennia multipuncta 0 0 1 1 0 0 0 1 0 1 1 0 0 1 0 0 0 0 1 0 1 0 1 1 1 1 1 1 0 – 1 0 0 0 1 0 0 0 030 Herennia papuana 0 0 1 1 0 0 0 1 0 1 1 0 0 1 0 0 0 0 1 0 1 0 1 1 1 1 1 1 0 – 1 0 0 0 1 0 0 0 031 Nephilengys malabarensis 1 1 0 0 0 0 1 0 0 1 1 1 0 1 0 0 0 0 1 0 0 0 1 1 1 1 1 0 0 – 1 0 0 0 1 0 0 0 032 Nephilengys cruentata 1 1 0 1 0 0 1 0 0 1 1 1 0 1 0 0 0 0 1 0 0 0 1 1 1 1 1 1 0 – 1 0 0 0 1 0 0 0 0

Characters

Taxa 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76

1 Deinopis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 – 0 0 0 0 0 0 0 – – – – – 0 0 0 – 0 02 Uloborus 0 0 0 1 1 0 0 0 0 0 0 0 1 0 0 1 0 0 – 0 0 0 1 0 0 0 – – – – – 0 0 0 – 1 03 Araneus 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 1 0 1 – 1 1 0 0 0 – – – – – 1 0 1 4 0 04 Argiope aurantia 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 1 1 1 0 0 0 1 0 1 0 0 – – – – – 1 0 1 4 0 15 Argiope argentata 0 0 0 0 0 0 1 0 0 0 0 0 1 0 1 1 1 1 0 0 0 1 0 1 0 0 – – – – – 1 0 1 4 0 16 Linyphia 0 0 0 0 0 0 0 1 0 1 1 – 1 0 0 0 0 0 – 1 – 1 1 0 0 0 – – – – – 0 0 0 – 1 07 Pimoa 0 0 0 0 0 0 0 1 0 1 1 – 1 0 0 0 0 0 – 1 – 1 1 0 0 0 – – – – – 0 0 0 – 0 18 Steatoda – – – 0 0 0 0 0 1 1 1 – 1 0 0 0 0 1 0 1 – 1 1 0 0 0 – – – – – 0 0 0 – 1 09 Nesticus – – – 0 0 0 0 0 1 1 1 – 1 0 0 0 0 0 – 1 – 0 1 0 0 0 – – – – – 0 0 0 – 1 010 Epeirotypus 0 0 0 0 0 0 0 0 0 0 1 – 1 0 0 0 0 1 0 1 – 1 1 1 0 0 – – – – – 0 1 0 – 1 011 Tetragnatha 0 0 0 1 0 0 0 0 0 0 1 – 0 1 0 0 0 1 0 1 – 1 1 0 0 0 – – – – – 1 1 1 4 1 012 Meta 0 0 0 0 0 0 0 0 0 0 1 – 1 0 0 0 0 1 0 0 0 1 1 0 0 0 – – – – – 0 0 1 4 0 013 Leucauge 0 0 0 1 0 0 0 0 0 0 1 – 1 0 0 0 0 0 – 1 – 1 1 0 0 0 – – – – – 1 1 1 0 1 014 Deliochus 0 0 0 0 0 0 0 0 0 1 1 – 1 0 0 0 0 ? 0 ? 0 1 1 0 0 0 – – – – – 0 0 0 – 1 015 Phonognatha graeffei 0 0 0 0 0 0 0 0 0 1 1 – 1 0 0 0 0 1 0 0 0 1 1 0 0 0 – – – – – 0 0 0 – 0 016 Clitaetra episinoides 0 0 0 0 0 0 0 0 0 0 0 1 1 3 0 0 0 1 1 0 0 1 1 0 0 0 1 0 0 1 0 1 0 1 0 1 017 Clitaetra perroti 0 0 0 0 0 0 0 0 0 0 0 1 1 3 0 0 0 1 1 0 0 1 1 0 0 0 1 0 0 1 0 1 0 1 0 1 018 Clitaetra clathrata 0 0 1 0 0 0 0 0 0 0 1 – 1 3 0 0 0 1 1 0 0 1 1 0 0 0 1 0 0 1 1 1 0 1 0 0 019 Clitaetra irenae 0 0 1 0 0 0 0 0 0 0 0 1 1 3 0 0 1 1 1 0 0 1 1 0 0 0 1 0 0 1 0 1 0 1 0 0 020 Clitaetra simoni 0 0 1 0 0 0 0 0 0 0 ? ? 1 3 0 0 0 1 1 0 0 1 1 1 0 0 1 0 0 1 1 1 0 1 0 1 021 Clitaetra thisbe 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 0 0 1 1 0 0 0 1 0 0 0 0 0 0 1 0 1 022 Nephila clavipes 0 0 0 0 1 1 1 0 0 0 0 1 1 1 0 0 0 1 0 0 0 1 0 1 0 0 1 0 1 0 0 1 0 1 3 0 123 Nephila fenestrata 0 0 0 0 1 1 1 0 0 0 0 1 ? ? 0 0 0 1 0 0 1 1 0 1 0 0 1 0 1 0 0 0 0 1 1 0 124 Nephila pilipes 1 0 0 0 1 1 1 0 0 0 0 1 0 1 0 0 0 1 0 0 1 1 0 1 0 0 1 0 0 0 0 0 0 1 3 0 125 Nephila antipodiana 0 1 0 0 1 1 1 0 0 0 0 1 0 1 0 0 0 1 0 0 1 1 0 1 0 0 1 0 1 0 0 1 0 1 1 0 126 Nephila clavata 0 1 0 0 1 1 1 0 0 0 0 1 1 0 0 0 0 1 0 0 0 1 0 1 0 0 1 0 1 0 0 1 0 1 1 0 127 Nephila plumipes 0 0 0 0 1 1 1 0 0 0 0 1 1 0 0 0 0 1 0 0 ? 1 0 1 0 0 0 – – – – 1 0 1 ? 0 128 Nephila edulis 0 0 0 0 1 1 1 0 0 0 0 1 1 0 0 0 0 1 0 0 0 1 0 1 0 0 0 – – – – 0 0 1 1 0 129 Herennia multipuncta 1 0 0 0 0 0 0 0 0 0 0 1 2 0 1 0 1 1 0 0 1 1 0 1 1 0 1 1 0 0 0 1 0 1 0 0 130 Herennia papuana 1 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 1 1 0 0 0 1 0 1 1 0 1 1 0 0 0 1 0 1 0 0 131 Nephilengys malabarensis 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 1 0 1 0 1 0 – – – – 1 0 1 2 0 132 Nephilengys cruentata 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 0 1 0 1 0 1 0 – – – – 1 0 1 2 0 1



Characters

Taxa 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110

1 Deinopis 0 0 0 0 – 0 – 0 0 0 0 0 0 0 0 0 0 0 0 0 – 0 0 0 0 – – – – 0 0 0 0 02 Uloborus 0 0 0 0 – 0 – 0 1 0 0 0 0 0 1 1 1 1 0 0 – 0 0 0 0 0 0 – 0 0 0 0 0 13 Araneus 0 0 0 0 – 0 – 1 1 1 0 0 1 – 1 ? 1 1 ? 1 0 1 0 1 0 0 0 1 0 0 0 1 0 14 Argiope aurantia 0 0 0 0 – 0 – 0 1 1 0 0 0 1 1 2 2 1 1 1 0 1 0 1 0 0 0 1 0 1 0 1 1 15 Argiope argentata 0 0 0 1 0 1 2 0 1 1 0 0 1 – 1 2 2 1 0 1 0 1 0 1 0 0 0 1 0 1 1 1 0 16 Linyphia 0 0 0 1 0 1 0 0 1 0 0 0 1 – 1 1 2 0 0 0 – 1 1 0 1 1 1 0 0 0 0 1 1 07 Pimoa 0 0 0 0 – 0 – 0 0 0 0 0 1 – 1 1 1 1 0 0 – 1 1 0 1 1 1 0 0 0 0 1 1 08 Steatoda 0 0 0 0 – 0 – 0 1 0 0 0 1 – 1 1 1 1 0 0 – 1 1 0 1 0 1 0 1 0 0 1 1 09 Nesticus 0 0 0 0 – 0 – 0 1 0 0 0 0 0 1 1 1 1 0 0 – 1 1 1 1 0 1 0 1 0 0 1 1 010 Epeirotypus 0 0 0 0 – 0 – 0 1 0 0 0 0 0 0 1 1 1 0 0 – 1 1 1 1 0 1 0 0 0 0 1 1 011 Tetragnatha – 0 0 – – – – – 0 0 – – 0 ? 1 – 0 0 0 0 – 1 0 1 1 0 1 0 0 0 0 0 1 012 Meta 0 1 0 0 – 0 – 0 1 0 0 0 1 – 1 1 2 0 0 0 – 1 0 1 1 0 1 0 0 0 0 1 1 013 Leucauge 0 0 0 0 – 0 – 0 1 0 0 0 1 – 0 1 2 1 0 1 0 1 0 1 1 0 1 1 0 0 0 1 1 014 Deliochus 0 0 0 1 0 1 1 0 1 0 0 0 1 – 1 1 0 1 0 1 0 1 0 1 1 0 1 0 0 ? 0 1 1 015 Phonognatha graeffei 1 0 0 1 0 ? 0 0 1 0 0 0 1 – 1 2 0 1 0 0 – 1 0 1 1 0 1 0 0 ? 0 1 1 016 Clitaetra episinoides 0 0 1 0 – 0 – 0 1 0 0 0 0 0 1 1 1 1 0 0 – 1 0 1 1 0 1 1 0 0 1 0 1 017 Clitaetra perroti 0 0 1 0 – 0 – 0 1 0 0 0 0 0 1 1 1 1 0 0 – 1 0 1 1 0 1 0 0 0 1 0 1 018 Clitaetra clathrata 0 1 0 0 – 0 – 0 1 0 1 1 0 0 1 1 1 1 0 0 – 1 0 1 1 0 1 1 0 ? 1 0 1 019 Clitaetra irenae 0 1 0 0 – 0 – 0 1 0 1 1 0 1 1 1 1 1 0 0 – 1 0 1 1 0 1 1 0 ? 1 0 1 020 Clitaetra simoni 0 1 0 0 – 0 – 0 1 0 1 1 0 0 1 ? ? ? ? 0 – 1 0 1 1 0 1 1 0 ? ? ? ? ?21 Clitaetra thisbe 0 0 0 0 – 0 – 0 1 0 0 0 0 0 1 1 1 1 ? 0 – 1 ? ? 1 ? ? ? ? ? ? ? ? ?22 Nephila clavipes 0 1 0 0 – 0 – 0 0 1 0 0 0 0 0 0 1 1 0 1 0 1 0 1 1 0 1 1 0 1 1 1 0 123 Nephila fenestrata 0 1 0 1 0 1 1 1 0 1 0 0 1 – 1 1 1 1 1 1 1 1 0 1 1 0 1 1 0 1 1 1 0 124 Nephila pilipes 1 1 0 0 – 0 – 0 0 1 0 0 0 0 0 1 1 1 0 1 1 1 0 1 1 0 1 1 0 1 1 1 0 125 Nephila antipodiana 1 1 0 0 – 0 – 0 0 1 0 0 0 0 0 1 1 1 0 0 – 1 0 1 1 0 1 1 0 1 1 1 0 126 Nephila clavata 1 1 0 0 – 0 – 0 0 1 0 0 1 – 0 1 1 1 0 0 – 1 0 1 1 0 1 1 0 1 1 1 0 127 Nephila plumipes ? 1 0 0 – 0 – 0 0 1 0 0 0 0 1 1 1 1 0 1 1 1 0 1 1 0 1 1 0 1 1 1 0 128 Nephila edulis 0 1 0 0 – ? 1 0 0 1 0 0 0 1 0 1 1 1 0 0 – 1 0 1 1 0 1 1 0 1 1 1 0 129 Herennia multipuncta 0 0 0 1 1 1 0 0 0 1 0 0 1 – 1 1 2 1 1 1 1 1 0 1 1 0 1 1 0 1 1 0 0 130 Herennia papuana 0 0 0 1 1 1 0 0 0 1 0 0 1 – 1 ? ? ? ? 1 1 1 0 1 1 0 1 1 0 1 1 0 0 131 Nephilengys malabarensis 0 0 0 1 0 1 1 0 1 1 0 0 1 – 1 1 1 1 1 0 – 1 0 1 1 0 1 1 0 1 1 0 0 132 Nephilengys cruentata 1 0 0 1 1 0 – 1 0 1 0 0 1 – 1 2 1 1 0 1 1 1 0 1 1 0 1 1 0 1 1 0 0 1

Characters

Taxa 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139

1 Deinopis 0 0 0 0 0 0 0 0 0 0 0 0 – – – – – – 0 0 – 0 0 – – 0 0 0 02 Uloborus 0 0 0 0 0 1 1 0 0 1 0 0 – – – – – – 0 0 – 0 1 0 0 0 0 0 03 Araneus 2 0 1 0 0 1 2 0 0 1 1 1 0 0 0 0 0 0 1 0 – 0 1 0 0 0 0 0 04 Argiope aurantia 0 0 0 1 0 0 1 0 0 1 1 1 0 0 0 0 0 0 1 0 – 0 1 0 1 0 0 0 05 Argiope argentata 2 0 0 0 0 0 1 0 0 1 1 1 0 0 0 0 0 0 1 0 – 1 1 0 1 0 0 0 06 Linyphia 1 0 0 0 0 0 1 0 0 1 1 1 – 3 1 0 0 0 0 0 – 0 0 – – 0 0 0 07 Pimoa 0 0 0 0 0 0 1 0 0 1 1 1 0 4 1 0 0 0 0 1 0 0 1 0 0 0 0 0 08 Steatoda 0 0 2 0 0 0 0 0 0 1 0 0 – – – – – – 0 0 – 0 1 1 0 0 0 0 19 Nesticus 0 1 2 0 0 0 1 0 0 1 1 1 0 3 1 1 0 0 0 1 0 0 1 1 0 0 0 0 110 Epeirotypus 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 ? 0 1 0 0 0 0 0 011 Tetragnatha ? 1 0 1 1 0 1 1 1 1 0 1 – 1 1 1 0 0 0 0 – 1 0 – – 0 0 0 012 Meta 0 0 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 1 1 0 0 0 – – 0 0 0 013 Leucauge 0 0 0 0 1 0 1 1 0 1 0 1 0 1 1 0 0 0 0 1 0 0 0 – – 0 0 0 014 Deliochus 0 0 0 0 0 0 1 0 0 1 1 ? – – 0 – – – 1 0 – 1 0 – – 0 0 0 015 Phonognatha graeffei 1 1 0 1 0 0 1 1 1 1 1 1 0 5 0 0 0 0 0 0 – 0 0 – – 0 0 0 016 Clitaetra episinoides 2 ? 0 ? 0 0 1 0 0 1 ? 1 1 2 1 0 1 1 0 1 0 0 0 – – 0 1 0 017 Clitaetra perroti 2 0 0 0 0 0 1 0 0 1 0 1 1 2 1 0 1 1 0 1 0 0 0 – – 0 1 0 018 Clitaetra clathrata 2 ? 0 ? 0 0 1 0 0 1 ? 1 1 2 1 0 1 1 0 1 0 0 0 – – 0 1 0 019 Clitaetra irenae 2 ? 0 ? 0 0 1 0 0 1 ? 1 1 2 1 0 1 1 0 1 0 0 0 – – 0 1 0 020 Clitaetra simoni ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?21 Clitaetra thisbe ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?22 Nephila clavipes 2 0 0 0 0 0 2 0 0 1 1 1 1 2 1 0 1 1 0 1 1 0 0 – – 0 0 0 023 Nephila fenestrata 2 0 0 0 0 0 1 0 0 1 1 1 1 2 1 0 1 1 0 1 1 0 0 – – 0 0 0 024 Nephila pilipes 2 0 0 0 0 0 1 0 0 1 0 1 1 2 1 0 1 1 0 1 1 0 0 – – 0 0 0 025 Nephila antipodiana 2 0 0 0 0 0 ? 0 0 1 1 1 1 2 1 0 1 1 0 1 1 0 0 – – 0 0 0 026 Nephila clavata 2 0 0 0 0 0 2 0 0 1 1 1 1 2 1 0 1 1 0 1 1 0 0 – – 0 0 0 027 Nephila plumipes 2 0 0 0 0 0 2 0 0 1 1 1 1 2 1 0 1 1 0 1 ? 0 0 – – 0 0 0 028 Nephila edulis 2 0 0 0 0 0 1 0 0 1 1 1 1 2 1 0 1 1 0 1 ? 0 0 – – 0 0 0 029 Herennia multipuncta 2 0 0 0 0 0 1 0 0 1 1 1 1 2 0 0 1 0 0 1 0 0 0 – – 1 0 0 030 Herennia papuana 2 0 0 0 0 0 1 0 0 1 1 1 1 2 1 0 1 0 0 1 0 0 0 – – 1 0 0 031 Nephilengys malabarensis 2 0 0 0 0 0 2 0 0 1 1 1 1 2 1 0 1 1 0 1 1 0 0 – – 0 0 1 032 Nephilengys cruentata 2 0 0 0 0 0 2 0 0 1 1 1 1 2 1 0 1 1 0 1 1 0 0 – – 0 0 1 0



Characters

Taxa 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168

1 Deinopis 0 0 0 0 0 – – – – – – – – 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 ? 02 Uloborus 0 0 0 0 0 – – – – – – – – 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 03 Araneus 0 0 0 0 0 – – – – – – – – 2 1 0 1 1 0 0 1 1 0 0 1 1 0 1 04 Argiope aurantia 0 1 0 0 0 – – – – – – – – 1 1 0 1 1 0 0 1 1 1 1 1 0 0 1 05 Argiope argentata 0 1 1 0 0 – – – – – – – – 1 1 0 1 1 0 0 1 1 1 1 1 1 0 1 06 Linyphia 1 – – – 0 – – – – – – – – 1 0 0 1 1 0 0 0 0 0 0 1 1 1 – –7 Pimoa 0 0 0 0 0 – – – – – – – – 1 0 0 0 0 0 0 0 0 0 0 1 1 1 – –8 Steatoda 0 1 1 0 0 – – – – – – – – 1 0 0 1 1 0 0 0 0 0 0 1 1 2 – –9 Nesticus 1 – – – 0 – – – – – – – – 1 0 0 0 0 0 0 0 0 0 0 1 1 2 – –10 Epeirotypus 0 1 0 0 0 – – – – – – – – 1 1 0 0 0 0 0 0 0 0 0 1 0 0 1 011 Tetragnatha 0 1 1 2 0 – – – – – – – – 1 0 0 1 1 0 0 0 0 0 0 1 1 0 0 012 Meta 0 1 1 0 0 – – – – – – – – 2 1 0 1 1 1 1 0 0 0 0 0 0 0 1 013 Leucauge 0 1 1 1 0 – – – – – – – – 1 0 0 1 1 0 0 0 0 0 0 1 1 0 ? 014 Deliochus 0 0 0 0 1 1 0 – 1 0 0 0 0 1 0 0 1 1 0 0 0 0 0 1 1 1 0 1 015 Phonognatha graeffei 1 – – – 1 1 0 – 1 0 0 1 0 1 2 0 1 1 0 0 1 0 0 0 1 1 0 1 016 Clitaetra episinoides 1 – – – 1 0 1 1 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 1 1 0 1 ?17 Clitaetra perroti 1 – – – 1 0 1 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 1 1 ? ? ?18 Clitaetra clathrata 1 – – – 1 0 1 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 1 1 0 ? ?19 Clitaetra irenae 1 – – – 1 0 1 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 1 ? 0 1 120 Clitaetra simoni ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?21 Clitaetra thisbe ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?22 Nephila clavipes 1 – – – 1 0 1 1 0 0 2 0 0 0 2 1 1 1 0 1 0 0 0 0 1 1 0 1 023 Nephila fenestrata 1 – – – 1 1 1 0 0 0 1 1 1 0 0 1 1 1 0 1 0 0 0 0 1 1 0 1 024 Nephila pilipes 1 – – – 1 0 1 1 0 0 0 0 0 0 2 1 1 1 0 1 0 0 0 0 1 1 0 1 025 Nephila antipodiana 1 – – – 1 0 1 1 0 0 2 0 0 0 2 1 1 1 0 1 0 0 0 0 1 1 0 1 026 Nephila clavata 1 – – – 1 1 1 0 0 0 1 0 0 0 2 1 1 1 0 1 0 0 0 0 1 1 0 ? ?27 Nephila plumipes 1 – – – 1 0 1 1 0 0 2 0 1 0 2 1 1 1 0 1 0 0 0 0 1 1 0 1 028 Nephila edulis 1 – – – 1 0 1 1 0 0 2 0 0 0 2 1 1 1 0 1 0 0 0 0 1 1 0 1 029 Herennia multipuncta 1 – – – 1 1 0 – 0 1 1 0 0 0 0 1 1 1 0 1 0 0 0 1 1 ? 0 1 130 Herennia papuana 1 – – – 1 1 0 – 0 1 1 0 0 0 0 1 1 1 0 1 0 0 0 1 1 0 0 1 131 Nephilengys malabarensis 1 – – – 1 1 0 – 0 1 1 0 0 0 0 1 1 1 0 1 0 0 1 0 1 1 0 1 032 Nephilengys cruentata 1 – – – 1 1 0 – 0 1 1 0 0 0 0 1 1 1 0 1 0 0 0 0 1 0 0 1 0

Characters

Taxa 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197

1 Deinopis 0 0 0 0 0 0 – – – 0 0 0 0 0 0 – – 0 0 0 0 – – 0 0 0 0 0 –2 Uloborus 0 1 0 0 0 1 – 0 0 1 0 1 0 0 0 – 0 0 1 0 0 – ? 0 0 0 1 0 –3 Araneus 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 – 0 0 0 0 0 0 0 0 0 0 0 0 –4 Argiope aurantia 0 1 0 0 0 0 0 0 0 0 0 ? ? 0 0 – 0 0 0 1 0 0 0 0 1 ? 0 0 –5 Argiope argentata 0 1 ? 0 0 0 0 0 0 0 0 1 0 0 0 – ? 0 0 1 0 0 0 0 1 1 0 0 –6 Linyphia 0 0 1 – – – – – – – – – – – – – – – 0 0 1 – ? 0 0 – 1 0 –7 Pimoa 0 0 1 – – – – – – – – – – – – – – – 0 0 1 – ? 0 ? – 1 0 –8 Steatoda 0 0 1 – – – – – – – – – – – – – – – 0 0 0 1 ? 0 0 – 0 0 –9 Nesticus 0 0 1 – – – – – – – – – – – – – – – 0 0 0 1 0 0 0 – 0 0 –10 Epeirotypus 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 – 1 1 0 0 1 – ? 0 0 0 0 0 –11 Tetragnatha 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 – 1 1 1 0 1 – 1 0 0 0 0 0 –12 Meta 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 – 1 1 0 0 0 0 0 0 0 0 0 0 –13 Leucauge 0 0 ? 0 0 0 1 0 1 0 0 0 0 0 0 – 1 1 0 0 0 0 1 0 0 0 0 0 –14 Deliochus 0 0 0 0 2 0 1 0 0 ? ? 1 0 0 0 – ? 0 0 0 1 – ? 1 0 0 0 1 115 Phonognatha graeffei 0 0 1 0 1 1 – 0 0 ? 0 1 1 0 1 0 0 0 0 0 1 – ? 0 0 0 0 2 116 Clitaetra episinoides 0 0 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? 0 ? ? ? 0 –17 Clitaetra perroti ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ?18 Clitaetra clathrata ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? 0 –19 Clitaetra irenae 0 0 0 1 1 1 – 1 0 1 1 1 1 1 1 0 1 2 0 0 1 – ? 0 1 1 1 0 –20 Clitaetra simoni ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?21 Clitaetra thisbe ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?22 Nephila clavipes 1 1 1 0 1 1 – 0 0 1 1 1 1 0 1 1 0 2 0 0 1 – 0 0 1 0 1 0 –23 Nephila fenestrata 1 0 1 0 1 1 – 0 0 ? ? 1 1 0 1 1 ? 2 0 0 1 – 0 0 1 0 ? 0 –24 Nephila pilipes 1 1 1 0 1 1 – 0 0 1 0 1 1 0 1 1 1 2 0 0 1 – 0 0 1 1 1 0 –25 Nephila antipodiana ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? ? ? 0 ? ? ? 0 –26 Nephila clavata ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?27 Nephila plumipes 1 0 1 0 1 ? ? 0 0 ? ? 1 1 0 1 1 ? ? 0 0 ? ? ? 0 0 0 ? 0 –28 Nephila edulis 1 0 1 0 1 ? ? 0 0 ? ? 1 1 0 1 1 ? ? 0 0 ? ? ? 0 ? ? ? 0 –29 Herennia multipuncta 0 0 0 1 ? ? ? 1 ? ? ? ? ? ? ? ? ? ? 0 0 ? ? ? 1 ? ? ? 0 –30 Herennia papuana 0 0 0 1 1 1 – 1 0 1 1 ? ? 1 1 ? ? 2 0 0 1 – 0 1 0 0 1 0 –31 Nephilengys malabarensis 0 0 1 1 1 ? ? 1 0 ? ? 1 1 0 1 ? 1 ? 0 0 1 – 0 1 ? ? ? 1 032 Nephilengys cruentata 0 0 1 1 1 1 – 1 0 1 1 1 1 0 1 1 1 2 0 0 1 – 0 1 1 0 1 1 0

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