u n i ve r s i t y o f co pe n h ag e n

Koslashbenhavns Universitet

The dinoflagellate cyst genera Achomosphaera Evitt 1963 and Spiniferites Mantell 1850in Pliocene to modern sedimentsMertens Kenneth Neil Van Nieuwenhove Nicolas Gurdebeke Pieter R Aydin HilalBogus Kara Bringue Manuel Dale Barrie De Schepper Stijn de Vernal Anne EllegaardMarianne Grothe Arjen Gu Haifeng Head Martin J Heikkila Maija Limoges AudreyLondeix Laurent Louwye Stephen Marret Fabienne Masure Edwige Matsuoka KazumiMudie Peta J Penaud Aurelie Pospelova Vera Price Andrea Michelle Ribeiro SofiaRochon Andre Sangiorgi Francesca Schreck Michael Torres Vladimir Uzar SerdarVersteegh Gerard J M Warny Sophie Zonneveld KarinPublished inPalynology

DOI1010800191612220181465739

Publication date2018

Document versionPublishers PDF also known as Version of record

Citation for published version (APA)Mertens K N Van Nieuwenhove N Gurdebeke P R Aydin H Bogus K Bringue M Zonneveld K(2018) The dinoflagellate cyst genera Achomosphaera Evitt 1963 and Spiniferites Mantell 1850 in Pliocene tomodern sediments a summary of round table discussions Palynology 42(Suppl 1) 10-44httpsdoiorg1010800191612220181465739

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Palynology

ISSN 0191-6122 (Print) 1558-9188 (Online) Journal homepage httpwwwtandfonlinecomloitpal20

The dinoflagellate cyst genera Achomosphaera Evitt1963 and Spiniferites Mantell 1850 in Pliocene tomodern sediments a summary of round tablediscussions

Kenneth Neil Mertens Nicolas Van Nieuwenhove Pieter R GurdebekeHilal Aydin Kara Bogus Manuel Bringueacute Barrie Dale Stijn De SchepperAnne de Vernal Marianne Ellegaard Arjen Grothe Haifeng Gu Martin JHead Maija Heikkilauml Audrey Limoges Laurent Londeix Stephen LouwyeFabienne Marret Edwige Masure Kazumi Matsuoka Peta J Mudie AureacuteliePenaud Vera Pospelova Andrea Michelle Price Sofia Ribeiro Andreacute RochonFrancesca Sangiorgi Michael Schreck Vladimir Torres Serdar Uzar Gerard JM Versteegh Sophie Warny amp Karin Zonneveld

To cite this article Kenneth Neil Mertens Nicolas Van Nieuwenhove Pieter R Gurdebeke HilalAydin Kara Bogus Manuel Bringueacute Barrie Dale Stijn De Schepper Anne de Vernal MarianneEllegaard Arjen Grothe Haifeng Gu Martin J Head Maija Heikkilauml Audrey Limoges LaurentLondeix Stephen Louwye Fabienne Marret Edwige Masure Kazumi Matsuoka Peta J MudieAureacutelie Penaud Vera Pospelova Andrea Michelle Price Sofia Ribeiro Andreacute Rochon FrancescaSangiorgi Michael Schreck Vladimir Torres Serdar Uzar Gerard J M Versteegh SophieWarny amp Karin Zonneveld (2018) The dinoflagellate cyst genera Achomosphaera Evitt 1963 andSpiniferites Mantell 1850 in Pliocene to modern sediments a summary of round table discussionsPalynology 42sup1 10-44 DOI 1010800191612220181465739

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Published online 14 Dec 2018

The dinoflagellate cyst genera Achomosphaera Evitt 1963 and SpiniferitesMantell 1850 in Pliocene to modern sediments a summary of round tablediscussions

Kenneth Neil Mertensa Nicolas Van Nieuwenhovebpad Pieter R Gurdebekec Hilal Aydind Kara Boguse ManuelBringuefg Barrie Daleh Stijn De Schepperi Anne de Vernalj Marianne Ellegaardk Arjen Grothel Haifeng GumMartin J Headn Maija Heikkileuroao Audrey Limogespad Laurent Londeixq Stephen Louwyec Fabienne MarretrEdwige Masures Kazumi Matsuokat Peta J Mudieu Aurelie Penaudv Vera Pospelovag Andrea Michelle PricewSofia Ribeirop Andre Rochonx Francesca Sangiorgil Michael Schrecky Vladimir Torresz Serdar UzardGerard J M Versteeghaaab Sophie Warnyac and Karin Zonneveldab

aIfremer LER BO Station de Biologie Marine Place de la Croix Concarneau CEDEX France bDepartment of Geoscience Aarhus UniversityAarhus Denmark cDepartment of Geology Ghent University Ghent Belgium dBiology Department Faculty of Sciences and Arts CelalBayar University Manisa Turkey eInternational Ocean Discovery Program Texas AampM University College Station TX USA fSchool of theEarth Ocean and Environment University of South Carolina Columbia SC USA gSchool of Earth and Ocean Sciences University of VictoriaVictoria BC Canada hGeosciences Department University of Oslo Oslo Norway iUni Research Climate Bjerknes Centre for ClimateResearch Bergen Norway jDepartement des sciences de la Terre et de lrsquoAtmosphere Geotop Universite du Quebec a Montreal MontrealCanada kDepartment of Plant and Environmental Sciences University of Copenhagen Frederiksberg C Denmark lMarine Palynology andPaleoceanography Laboratory of Palaeobotany and Palynology Department of Earth Sciences Faculty of Geosciences Utrecht UniversityUtrecht The Netherlands mThird Institute of Oceanography SOA Xiamen China nDepartment of Earth Sciences Brock University StCatharines Ontario Canada oEnvironmental Change Research Unit Department of Environmental Sciences University of Helsinki FinlandpDepartment of Glaciology and Climate Geological Survey of Denmark and Greenland (GEUS) Copenhagen Denmark qUniversite deBordeaux UMR 5805 EPOC Pessac CEDEX France rSchool of Environmental Sciences University of Liverpool Liverpool UK sCR2P Centrede Recherche sur la Paleobiodiversite et les Paleoenvironnements UMR7207 ndash CNRS MNHN UPMC Paris France tInstitute for East ChinaSea Research (ECSER) Nagasaki Japan uGeological Survey of Canada Dartmouth Canada vUMR6538 Domaines Oceaniques IUEM-UBOPlouzane France wDepartment of Geography McGill University Montreal Canada xISMER-UQAR 310 allee des Ursulines Rimouski QCCanada yDepartment of Geology University of Tromsoslash ndash The Arctic University Tromsoslash Langnes Norway zBiostratigraphy Core GroupExxonMobil Exploration Company Spring Texas USA aaDepartment of Marine Geochemistry Alfred-Wegener-Institut (AWI) HelmholtzZentrum feurour Polar und Meeresforschung Bremerhaven Germany abMARUM ndash Center for Marine Environmental Sciences Bremen UniversityBremen Germany acDepartment of Geology and Geophysics and Museum of Natural Science Louisiana State University Baton Rouge LAUSA adDepartment of Earth Sciences University of New Brunswick Fredericton Canada

ABSTRACTWe present a summary of two round-table discussions held during two subsequent workshops inMontreal (Canada) on 16 April 2014 and Ostend (Belgium) on 8 July 2015 Five species of the genusAchomosphaera Evitt 1963 and 33 of the genus Spiniferites Mantell 1850 emend Sarjeant 1970 occuringin Pliocene to modern sediments are listed and briefly described along with remarks made by workshopparticipants In addition several holotypes and topotypes are reillustrated Three species previouslyassigned to Spiniferites are here consideredaccepted as belonging to other genera Impagidinium inae-qualis (Wall and Dale in Wall et al 1973) Londeix et al 2009 Spiniferites rubinus (Rossignol 1962 exRossignol 1964) Sarjeant 1970 and Thalassiphora balcanica Baltescedil 1971 This summary forms the basisfor a set of papers that follows where points raised during the workshops are explored in greater detail

KEYWORDSSpiniferites AchomosphaeraHafniasphaeraRottnestia Pterocysta

1 Introduction

This chapter summarises discussions on the dinoflagellatecyst genera Achomosphaera Evitt 1963 and SpiniferitesMantell 1850 emend Sarjeant 1970 held during two work-shops The first took place at GEOTOP Universite duQuebec a Montreal (UQAM) in Montreal (Canada) on 16

April 2014 During this workshop all Spiniferites andAchomosphaera species recorded in Quaternary depositswere discussed individually by the participants Severalissues were noted regarding their classificationdescrip-tion and suggestions were made as how to resolve suchissues Several of the problems were further consideredduring a follow-up workshop at the Flanders Marine

CONTACT Email kennethmertensifremerfrSupplemental data for this article can be accessed httpsdoiorg1010800191612220181465739

2018 The Author(s) Published by AASP ndash The Palynological SocietyThis is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (httpcreativecommonsorglicensesby-nc-nd40)which permits non-commercial re-use distribution and reproduction in any medium provided the original work is properly cited and is not altered transformed or built upon in any way

PALYNOLOGY2018 VOL 42 NO S1 10ndash44httpsdoiorg1010800191612220181465739

Institute (VLIZ) in Ostend (Belgium) where a round-tablediscussion was held on 8 July 2015 Notes made by NVand PG during the discussions were summarised after-wards by NV PG and KNM and form the basis ofthis document

The aim of the workshops was to evaluate the taxonomyand nomenclature of those taxa assigned to the generaAchomosphaera and Spiniferites that have been recorded inPliocene to modern sediment We have compiled all rele-vant information on these taxa and reillustrated selectedholotypes and topotypes A generic overview that includesother related genera can be found in Mertens amp Carbonell-Moor (2018) We have excluded some Paratethyan taxa (iespecies described from deposits of the Paratethys Sea not-ably in the Pannonian Vienna Molasse and Ponto-Caspianbasins) that according to PJM lack unambiguous age con-straints for a Pliocene to modern occurrence (egSpiniferites maisensis Seurouto 1994 and Spiniferites virgulaefor-mis Seurouto 1994) All taxa belonging to Achomosphaera orSpiniferites that are currently known from Pliocene to mod-ern sediments are listed alphabetically in Section 2 Speciespreviously assigned to Spiniferites or Achomosphaera butnow considered to belong to other genera are listedalphabetically in Section 3 Note that the views presentedhere reflect a broad consensus and do not necessarillyimply full agreement among the panelists Authorships forall taxonomic names cited are provided in SupplementaryAppendix 1 and 2 We have updated stratigraphic rangeswhere possible to allow for the lowering of the base of thePleistocene in 2009 from the base of the Calabrian Stageto that of the Gelasian Stage effectively from 18 to 26MaA consequence was the readjustment of the Piacenzianfrom Middle Pliocene which had become irrelevant toUpper Pliocene (Gibbard and Head 2010 Head andGibbard 2015)

2 Discussion of taxa belonging to the generaAchomosphaera and Spiniferites occurring inPliocene to modern sediments

ll taxa are listed alphabetically with their synonyms (withrelated comments in brackets) and other relevant informa-tion Distinguishing characters are described using existingliterature that has been paraphrased and updated withmodern terminology Intraspecific morphotypes here referto formal subspecies (or varieties) that have been describedor informal morphologies described in the literature Pointsfrom the discussions that arose during the workshops aredocumented in the remarks section for each speciesdescribed and elsewhere where appropriate Often noagreement could be reached to make any formal changesand this document then reflects this disagreement Forexample some participants preferred to use Spiniferites mul-tisphaerus while others considered this species to belongto Hafniasphaera such disagreements are left open in thissummary For wall structure we follow terminology used byHead (1994) Measurements are those given in cited refer-ences Kofoidean nomenclature is used to designate the

plate numbers For simplicity we choose not to use lsquopara-rsquoterminology to distinguish features of cysts from theirmotile counterparts since only cyst morphology isaddressed here

21 Achomosphaera andalousiensis Jan du Chene 1977emend Jan du Chene and Londeix 1988

Synonymy ldquoAchomosphaera perforatardquo Morzadec-Kerfourn1979 pl 31 figs 1ndash2 4 1984 pl II figs 13ndash14non Spiniferites septentrionalis Harland 1979 p 103ndash104 pl1 figs 12ndash18 text-fig 4Holotype Jan du Chene 1977 pl 1 fig 1 lost according toJan du Chene amp Londeix (1988 p 237)Lectotype Jan du Chene and Londeix (1988 p 244 pl 1fig 1ndash3)Type locality Carmona section Andalusia SpainType stratum Upper Miocene (Jan du Chene 1977)Etymology Derived from the type locality (Jan duChene 1977)Distinguishing characters Ellipsoid central body bearinglong slender processes The pedium is thin and smooththe tegillum is thick and smooth to shagreenate Pediumand tegillum are closely appressed except below the proc-esses which are formed by the tegillum The processes aregonal and long slender hollow and sometimes fenestrateat their bases and terminate distally in characteristic fenes-trate platforms Some adjacent processes (often in apical orcingular areas) are connected by crests The sulcus is indi-cated by small processes with bifid distal ends sutural orna-mentation is completely absent The archeopyle is formedby loss of plate 300 the operculum is free (Based on Jan duChene 1977 p 112 and Jan du Chene amp Londeix 1988p 239ndash241)Dimensions Central body length 40ndash50 mm width34ndash44 mm process length 14ndash26 mm width of distal ends14ndash21 mm (Jan du Chene amp Londeix 1988)Biological affinity UnknownIntraspecific morphotypes Achomosphaera andalousiensissubsp suttonensis Head 1997 has many fenestrations in thedistal platforms of the processes and was described from theLower Pliocene of eastern England (Head 1997)Achomosphaera andalousiensis subsp andalousiensis (auto-nym) has notably fewer fenestrations in the distal platformsof the processesComparison For Spiniferites septentrionalis see Section 235Spiniferites speetonensis from the Lower Cretaceous has simi-lar distal process endings but differs in having intergonalprocesses and sutures (Jan du Chene amp Londeix 1988)Spiniferites perforatus from the Lower Eocene differs fromAchomosphaera andalousiensis in having intergonal processesand sutures that are sometimes elevated (Jan du Chene ampLondeix 1988)Remarks Spiniferites septentrionalis is a junior synonymaccording to Harland (1983 p 103ndash104) whose opinion wasfollowed by Jan du Chene amp Londeix (1988 p 421) but ques-tioned by Head amp Wrenn (1992 p 2) and not accepted here

PALYNOLOGY 11

ldquoAchomosphaera perforatardquo of Morzadec-Kerfourn 1979from the Lower Holocene of Tunisia as remarked by LL andKNM it is unclear whether Morzadec-Kerfourn (1979) incor-rectly used this name to refer to Achomosphaera ramuliferavar perforata from the Lower Eocene (later transferred toAchomosphaera ramulifera subsp perforata) or intended topropose a new combination Either way Morzadec-Kerfourndid not validly publish the name Achomosphaera perforataMorzadec-Kerfourn (1979) synonymised Achomosphaeraandalousiensis and ldquoAchomosphaera septentrionalisrdquo withAchomosphaera ramulifera subsp perforata we do not agreewith this synonymy

There is some doubt about the stratigraphic range ofAchomosphaera andalousiensis (see eg Head 1996ap 1207) but if the specimens of Morzadec-Kerfourn (1979)from the Upper Pleistocene to Upper Holocene of Tunisiaare in situ the species ranges from the Serravallian

(Middle Miocene Dybkjaer amp Piasecki 2010) to theLate Holocene

22 Achomosphaera argesensis Demetrescu 1989(Plate 1 figures 1ndash6)

Synonymy NoneHolotype Demetrescu 1989 pl 1 figs 1ndash5 textndashfig 2 text-fig 3A CType locality No specific type locality was clearly specifiedby Demetrescu (1989) but it is presumably somewhere inthe Arges region of RomaniaType stratum Lower Pliocene of the southern Carpathiansforedeep (Demetrescu 1989)Etymology Named after the Arges region of Romania(Demetrescu 1989)

Plate 1 1ndash6 Achomosphaera argesensis Demetrescu 1989 from S4677 Hungary FI-2007-001 at high to low focus note hollow bases of processes (shown by bub-bles present in 2ndash3) visible tabulation (3) thick wall (4) and high crest above archeopyle (5) Slide provided by VTT photographed by KNM All scale bars frac14 10mm

12 K N MERTENS ET AL

Distinguishing characters Ovoid central body with a pro-nounced apical boss and lacking sutural ornamentation Theouter surface is smooth and bears 24 lobate branched exclu-sively gonal processes which can have fenestrate bases Theprocesses are hollow or sometimes partly solid Their distalends expand into lobate short branches and some have irregu-larly fringed tips Two of the apical processes are fused andhave a flattened distal end which can also be the case for twoof the antapical processes Expressed tabulation is 40 600 6cs5ndash6000 1p 100 00 The ends of the cingulum are displaced byapparently about one cingulum width and the sulcus isstraight The archeopyle is formed by loss of plate 3rdquo and theoperculum is free (Based on Demetrescu 1989 p 51ndash53)Dimensions Central body length 47ndash50 mm width 32ndash35mmthickness 33ndash35 mm apical boss 1ndash7 mm long process length15ndash26 mm (Demetrescu 1989)Biological affinity UnknownIntraspecific morphotypes NoneComparison Achomosphaera argesensis differs fromAchomosphaera ramulifera in having an apical boss whichresembles that of Achomosphaera improcera Islam 1983 fromthe Eocene and Spiniferites bentorii but Achomosphaeraimprocera has much shorter processes and Spiniferites ben-torii has low sutures Achomosphaera argesensis also resem-bles Achomosphaera andalousiensis but the latter does nothave a flat apical process or processes with lobate tipsSpiniferites validus Seurouto-Szentai 1982 described from thePannonian Basin differs from Achomosphaera argesensis inhaving a thin elongated apical process instead of a largeflattened apical process (Based on Demetrescu 1989)Remarks PJM was unsuccessful in locating the holotype inBucharest because the Institute of Geology Bucharest hasbeen dismantled and the location of the type slides is nolonger known moreover it was not possible to contactEmanuel Demetrescu so the holotype is effectively lostEaton (1996 pl IV figs 6ndash7) illustrated well-preserved speci-mens from the Black Sea with membranes connecting thedistal ends of apical and antapical gonal processes Eatonrsquosslides are archived in the Palynological Slide Collection ofthe Department of Paleontology at the Natural HistoryMuseum London UK The age of Eatonrsquos sample is uncer-tain VT noted that he saw specimens of Achomosphaeraargesensis from the Pannonian Basin However based on theoriginal publication of Demetrescu (1989) VT identifiedAchomosphaera argesensis on the basis that the holotypeillustrations show clear membranes connecting the gonalprocesses associated with the apical plates He suggestedthat this is the criterion to consider in separatingAchomosphaera argesensis from the other species describedfrom the Pannonian BasinParatethys AG agreed that it isvery distinct and suggested that these forms ofAchomosphaera may fit better in Spiniferites During the draft-ing of this report LL remarked that a lack of true septabetween most processes would still leave Achomosphaeraargesensis in Achomosphaera FS noted during drafting thatshe observed Achomosphaera argesensis in Upper Miocene(regional Pontian Stage) sediments of the ParatethyanBlack Sea

23 Achomosphaera callosa Matsuoka 1983 (Plate 2figures 1ndash3)

Synonymy NoneHolotype Matsuoka 1983 pl 11 figs 6andashcType locality Ota Tsugawa-cho Niigata Prefecture cen-tral JapanType stratum Tokonami Formation equivalent to theNishiyama Formation Pliocene (Matsuoka 1983)Etymology From Latin callosa thick skinned in reference tothe thick wall (Matsuoka 1983)Distinguishing characters Central body thick-walled spher-oidal to (rarely) ovoid with a coarsely granular outer surfaceThe smooth processes are gonal only and sutural septa areonly occasionally present The archeopyle is formed by lossof plate 3rdquo and is reduced (Based on Matsuoka 1983p 128ndash129)Dimensions Central body length 36ndash53 mm width 36ndash45 mmwall thickness up to 3 mm process length up to 15 mm(Matsuoka 1983 p 128)Biological affinity UnknownIntraspecific morphotypes NoneComparison Achomosphaera callosa resemblesAchomosphaera crassipellis (Deflandre amp Cookson 1955)Stover amp Evitt 1978 and Achomosphaera cf sagena Davey ampWilliams 1966 of Gocht (1969 p 36 pl 7 figs 1ndash2) but dif-fers from these two forms in having a coarsely granular outersurface and exclusively gonal processes (Matsuoka 1983) Thecentral body of Achomosphaera crassipellis is larger(74ndash89 mm) than that of Achomosphaera callosa and has athicker wall judging from the illustration in Deflandre ampCookson (1955 no measurements provided) and longer proc-esses (23ndash26 mm) (measurements from Deflandre ampCookson 1955)Remarks Matsuoka (1983 p 128ndash129) describedAchomosphaera callosa from the Pliocene and LowerPleistocene of Japan LL expressed that he has no problemwith the species although he has not observed it inQuaternary sediments The exact range of the speciesremains to be determined MJH remarked that the centralbody apparently has a solid pedium a prismatic layer on topand open luxuria but no tegillum This leads to the questionof what the processes are made of an issue probably bestanswered through scanning electron microscopy Other thanthat MJH considers Achomosphaera callosa a distinctive spe-cies and LL agrees

24 Achomosphaera granulata Mao Shaozhi 1989

Synonymy Non Achomosphaera sp A of Matsuoka 1983 pl11 figs 1ndash5 illustrated in Plate 2 figures 4ndash8Holotype Mao Shaozhi 1989 pl 28 fig 10Type locality Northern Xisha Trench South China SeaType stratum Upper Pleistocene (Mao Shaozhi 1989)Etymology Derived from the occurrence of dense granuleson the wall and processes (Mao Shaozhi 1989)Distinguishing characters Central body ovoid thick-walledlight brown to brown commonly with a short apical boss

PALYNOLOGY 13

The apical horn has a basal width greater than its length(typically 3ndash5 mm) and has a truncated distal end The cingu-lum is 5ndash7mm wide delimited by dense granules separatingthe cyst into a sub-triangular epicyst and rounded to nearlytrapezoidal hypocyst There are two wall layers the thick

outer wall forming the exclusively gonal processes The sur-face of the outer wall is ornamented by dense and well-dis-tributed granules The granules are coarse and sometimeslink to form curved lines They are developed on both thecentral body and processes and make the outline of the

Plate 2 1ndash3 Holotype of Achomosphaera callosa Matsuoka 1983 at high to low focus 4 Topotype 5 Focus on holotype showing detail of wall texture and pro-cess 6ndash8 Upper to lower focus of Achomosphaera sp A of Matsuoka (1983) plate 11 figures 1ndash5 [Mao Shaozhi (1989) considered her specimens ofAchomosphaera granulata the same as these specimens from the Lower to Upper Miocene of the Niigata district (central Japan) but we disagree with this assess-ment] Slides provided by KM photographed by KNM All scale bars frac14 10mm

14 K N MERTENS ET AL

processes rough The processes can be perforated due to cor-rosion according to Mao Shaozhi (1989) The processes havewide bases and taper rapidly No sutures are present exceptfor the granules that mark the cingulum dorsally The archeo-pyle is formed by loss of plate 3rdquo (Based on Mao Shaozhi1989 p 139 translated from Chinese by HG and observationsof Achomosphaera sp A of Matsuoka 1983 p 130)Dimensions Central body length (including apical horn)45ndash53 mm width 37ndash45 mm wall thickness 2 mm processlength 10ndash13 mm (Mao Shaozhi 1989)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species can be distinguished from all simi-lar Spiniferites and Achomosphaera species by the uniquedense granules covering the wall and processes its thickouter wall and brown colour (Mao Shaozhi 1989)Remarks Mao Shaozhi (1989) considered Achomosphaeragranulata the same as Achomosphaera sp A of Matsuoka1983 from the Lower to Upper Miocene of the Niigata dis-trict central Japan but we disagree with this assessmentsee also Londeix et al (2018)

The species has been recorded only in Quaternary sedi-ments of the South China Sea by Mao Shaozhi (1989) andMao Shaozhi amp Harland (1993) Londeix et al (2018) consid-ered this species to belong to Hafniasphaera others includ-ing KNM and VP were not convinced because the publishedholotype images are not sharp and the description does notmention the presence of vacuoles

25 Achomosphaera ramosasimilis (Yun Hyesu 1981)Londeix et al 1999

Synonymy Achomosphaera ramulifera subsp ramosasimilisYun Hyesu 1981 p 14ndash15 pl 1 figs 1 8 text-fig 3b

Spiniferites ramuliferus (auct non Deflandre) Reid Reid1974 p 608 pl 4 figs 39ndash40Holotype Yun Hyesu 1981 pl 1 fig 1 text-fig 3b Fensomeet al 1991 figs 1ndash2 p 719 fig 4 p 719 amp 721 [initiallydescribed in German by Yun Hyesu 1981 translated byFensome et al 1991]Type locality Timmermann brickyard new pit near EsbeckWestphalia GermanyType stratum Santonian Upper Cretaceous (Yun Hyesu 1981)Etymology In reference to the close similarity between thisspecies and Spiniferites ramosus (Fensome et al 1991 p 721)Distinguishing characters Thick-walled ovoid to spheroidalcentral body with a granular or smooth pedium and asmooth tegillum bearing hollow distally closed exclusivelygonal processes The processes on the cingulum may be con-nected by crests similar to some processes in the apical andantapical areas There is always a thin apical process occa-sionally with a terminal hooklet Sutures can be partly pre-sent The archeopyle is formed by the loss of plate 3rdquo (Basedon Yun Hyesu 1981 p 14ndash15) and Fensome et al 1991p 719ndash720)Dimensions Central body length 30 (32) 36 mm width 32(39) 42 mm maximum length of processes 16ndash18 mm (YunHyesu 1981)

Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Yun Hyesu (1981) becauseAchomosphaera ramosasimilis possesses a thick wall partialpresence of sutures does not have all processes connectedby crests bears a simple apical process and no antapicalbranched process it differs from Achomosphaera ramuliferaRemarks This species was observed by Londeix et al (1999)from the Lower Pliocene of the Strait of Sicily centralMediterranean Sea LL also considers specimens ofldquoSpiniferites ramuliferusrdquo described by Reid (1974) from recentraised-beach deposits at Woodgrange Northern Ireland UKto be probably Achomosphaera ramosasimilis and notAchomosphaera ramulifera see discussion in Londeix et al(2018) and Gurdebeke et al (2018)

26 Spiniferites alaskensis Marret et al 2001 ex Marretin Fensome amp Williams 2004

Synonymy NoneHolotype Marret et al 2001 pl 1 figs 7ndash9 designated byMarret in Fensome amp Williams (2004)Type locality ODP Site 887 Gulf of Alaska northeasternNorth Pacific Gulf of AlaskaType stratum Marine Isotope Substage 5e Sample 887B-2H-5 65 cm (Marret et al 2001)Etymology Named alaskensis from the type locality (Marretet al 2001)Distinguishing characters Ovoid shape with an apical bossThe cyst wall is thin (lt 1 mm) and has a finely granulate toscabrate surface Processes are gonal and relatively broad(2 mm) each process terminating in a simple trifurcation withpointed ends Low sutural septa are present between proc-esses and define a gonyaulacacean tabulation The archeo-pyle is formed by loss of plate 3rdquo (Based on Marret et al2001 p 384ndash385)Dimensions Central body length 263 (314) 368 mm width236 (293) 315 mm process length 75 (101) 125 mm (Marretet al 2001)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species differs from other Spiniferites spe-cies only by the process shape and termination Spiniferitesalaskensis has broad processes with simple short trifurcatebranches with pointed ends such processes are unusual forlate Quaternary Spiniferites species (Marret et al 2001p 386)Remarks The name was validated by Marret in Fensome ampWilliams (2004) because Marret et al (2001) did not indicatewhich of the illustrations represented the holotype This spe-cies is discussed in Marret amp Mertens (2018)

27 Spiniferites asperulus Matsuoka 1983 (Plate 3figures 1ndash6)

Synonymy NoneHolotype Matsuoka 1983 pl 12 fig 2

PALYNOLOGY 15

Plate 3 1ndash3 Holotype of Spiniferites asperulus Matsuoka 1983 in ventral view at high to low focus 4 Same specimen mid-focus showing wall structure 5 Samespecimen mid-focus showing antapical processes 6ndash8 Holotype of Spiniferites firmus Matsuoka 1983 in ventral view at high to low focus 9 Same specimen show-ing microgranular wall 10 Same specimen showing distal ends of cingular processes Slides provided by KM photographed by KNM All scale bars frac14 10 mmexcept 1ndash3 7ndash9frac14 20 mm

16 K N MERTENS ET AL

Type locality Nishiyama Nishiyama-cho Niigata PrefectureCentral JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin asper (somewhat rough) in ref-erence to the coarsely granular surface of the cyst (Matsuoka1983 p 131)Distinguishing characters The subspheroidal to ovoid cysthas a relatively thick wall The tegillum forms many smallinvaginations in the intratabular area No granules are pre-sent The processes are slender membranous gonal (andoccasionally intergonal) and widen at the base the twoantapical processes are somewhat more robust and con-nected by a slightly elevated crest (Based on Matsuoka1983 p 131ndash132 and observations by KNM)Dimensions Central body length 48ndash69 mm width45ndash64 mm process length up to 16 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison The wall structure is similar to that ofSpiniferites ludhamensis but there are many more invagina-tions in Spiniferites asperulus and Spiniferites ludhamensis issmaller (central body length 38ndash49 mm) and has completelyhollow processes Spiniferites membranaceus has a granu-lar wallRemarks LL expressed the difficulty in differentiating thisspecies from others such as Spiniferites firmus KM stated thatafter having described it back in 1983 he now has no clearrecollection of what this species is LL looked at the holotypebut adds that he did not look at the total population andtherefore finds it hard to say if this is a distinct species or notHe noted that granulation which may occur as a variation inmany species could be a response to specific environmentalconditions The original description mentions the occasionaloccurrence of intergonal processes a feature confirmed byKM This species is discussed in Limoges et al (2018)

28 Spiniferites belerius Reid 1974

Synonymy Non Spiniferites belerius sensu Harland 1977 p98ndash99 pl 1 figs 7ndash10 pl 2 figs 7ndash10 16ndash21 25ndash57 [frac14Spiniferites membranaceus]Holotype Reid 1974 pl 2 figs 12ndash13Type locality Severn Estuary England UKType stratum Modern surface sediments (Reid 1974)Etymology From the Latin belerium which is the Romanname for the promontory on which Landrsquos End CornwallEngland is situated (Reid 1974 p 596)Distinguishing characters The small central body is ovalwith a finely granular wall and an apical boss The processes aregonal and connected by low crests with a typical process shapeThis species was described as having a characteristic largelsquotrumpet-shapedrsquo posterior process at the junction of plates 1and 2 (Reid 1974 p 597 but see Gurdebeke et al 2018)Dimensions Central body length 35ndash42 mm width28ndash37 mm maximum process length 7ndash10 mm posterior pro-cess length 10ndash15 mm (Reid 1974)

Biological affinity Gonyaulax scrippsae Kofoid 1911 asdepicted by Wall amp Dale (1968 p 270 pl 1 fig 14) accord-ing to Reid (1974) LL and KZ noted some similarity to thecyst of Gonyaulax baltica Ellegaard Lewis amp Harding 2002 asproduced in batch-cultures and illustrated by Ellegaard et al(2002) KZ felt she could not understand the cyst-based spe-cies concept here because of the large morphological vari-ation documented in that study VP wondered if anyonecould distinguish cysts of Gonyaulax baltica in the sedimentsME replied that some cysts of Gonyaulax baltica could be thesame as Spiniferites belerius but ideally we should first deriveSpiniferites belerius in culture and compare the phylogeneticsIntraspecific morphotypes NoneComparison LL noted during the first workshop thatSpiniferites bentorii is larger overall and has a hypocyst largerthan the epicyst Spiniferites belerius is similar to Spiniferitesconiconcavus but the latter has wider process basesRemarks During the first and second workshops there wasagreement that this species is problematic LL consideredthis species as having a large morphological variation Healso suggested that this species is defined by central bodysize and shape and the presence of short processes that arenot well developed in the apical region PJM mentioned thatspecimens from the Black Sea show a very wide morpho-logical range and that these could be used as an illustrationof the speciesrsquo various morphologies KNM stated howeverthat potentially different species have been described asSpiniferites belerius given the great variation that he hasobserved globally PJM remarked that this is hard to tellwithout culture studies LL proposed that two categories bedefined for practical purposes typical and atypical morpho-types These categories can then be used in paleoecologicalstudies LL further underlined the wide range in morphologythat appears to characterise many Paratethyan species PJMasked LL whether he considered Spiniferites belerius to beParatethyan and LL answered ldquonordquo He noted that there is aclear regular shape and a wide range of morphological vari-ability and that it is sometimes hard to know if this repre-sents infraspecific variability or more than one species WhenPJM asked what LL considers a Paratethyan species hereplied that it is an ldquoendemicrdquo species with a clearly definedtypical morphology He used Seriliodinium explicatum Eaton1996 Invertocysta sp A of Londeix et al (2009)Invertocysta sp B of Londeix et al (2009) Pterocysta crucifor-mis Rochon et al 2003 Pyxidinopsis psilata (Wall amp Dale inWall et al 1973) Head 1994 Galeacysta etrusca Corradini ampBiffi 1988 and Caspidinium rugosum Marret in Marret et al2004 as good examples of Paratethyan species PJM com-mented in draft that LLrsquos examples include taxa not recordedoutside the Ponto-Caspian basins together with taxa havingwide ranges from the saline western Mediterranean Sea tothe brackish Aral Sea She considers it difficult to justify theword endemic for such a broad grouping of taxa She con-siders that by definition an endemic species means native orrestricted to a certain country or area and should not beused for Paratethyan forms as the Paratethys was a vastEuro-Asian waterway BD suggested that we should decidewhether we can put species into two different groups those

PALYNOLOGY 17

that are stable and well-described and those that originatefrom particular extreme environments that induce increasedecological pressure He said that much of the discussionappears to be focused on forms from extreme environmentsHe noted however that he does not know how to applysuch an approach in practice but proposed that ldquoweirdrdquomorphotypes should be addressed ldquolater onrdquo once clearbasic forms have been established KNM suggested that thetopotype material of Spiniferites belerius should be looked atto evaluate the morphological variability present This hasnow been done by Gurdebeke et al (2018) Limoges et al(2018) have accepted the presence of additional intergonalprocesses in this species

29 Spiniferites bentorii (Rossignol 1964) Wall ampDale 1970

Synonymy Hystrichosphaera bentorii Rossignol 1964 p84ndash85 pl 1 figs 3 3 inset 5ndash8 pl 3 figs 2ndash3 text-figs AndashF

Leptodinium churchillii Harland 1968 p 548 550ndash551 figs12ndash13 22ndash24

non Spiniferites nodosus (Wall 1967) Sarjeant 1970[Synonymy proposed by Reid (1974 p 599) is here rejected]

non Spiniferites bentorii sensu Wall amp Dale 1970 p 52 pl1 figs 26 28Holotype Rossignol 1964 pl 1 figs 3 7ndash8Type locality Ashdod borehole coastal plain IsraelType stratum Quaternary sediments 172ndash1725m depth(Rossignol 1964)Etymology Named after Dr Yaakov Ben-Tor (b 1910 ndash d2002) Israeli geologist who was head of the IsraeliGeological Survey at the time of Martine Rossignolrsquos studyand provided her with samplesDistinguishing characters This species typically has a pear-shaped central body with a characteristic apical boss It bearscharacteristic tapering slender gonal and occasionally inter-gonal processes with the two antapical processes being thelongest The process bases may be fenestrate Sutures aremarked by low ridges and sometimes vacuoles are presentin the sutures The wall is smooth to microgranular The cin-gular displacement is relatively large (three times its widthaccording to Reid 1974 one and a half to two times itswidth according to Wall 1965) Tabulation is typical for thegenus according to Wall (1965) and Harland (1968) with fourapical plates although the suture between 1 and 2 is faintand was not observed by Rossignol (1964) or Price ampPospelova (2014) both interpreting this as indicating thepresence of only three apical plates visible on the cyst Thesulcus is often well expressed and straight and widens poster-iorly (Reid 1974) The archeopyle is formed by loss of plate 3rdquoand is reduced with rounded corners (Based on Rossignol1964 p 84ndash85 Harland 1968 p 542ndash543 Reid 1974p 598ndash600 Rochon et al 1999 p 34 Price amp Pospelova2014 p 13 and observations of the participants) MJH notedin draft that specimens from the Last Interglacial of theBaltic have distinctive lateral cingular processes in which thedistal furcations branch abruptly at 90 to the process shafts

and point towards both poles (Head 2007 fig 7jkm)MJH further noted in draft that specimens of S bentorii thathe has observed have archeopyles well defined bythe suturesDimensions Central body length 60ndash73 mm width45ndash63 mm process length 15ndash20 mm (25 mm for the antapi-cal processes) (Rossignol 1964) Central body length58ndash69 mm width 52ndash55 mm length of processes 0ndash20 mm(Reid 1974)Biological affinity Gonyaulax digitale (Pouchet 1883) Kofoid1911 according to Wall amp Dale (1967 p 352) and Dodge(1989 p 283)Intraspecific morphotypes Rossignol (1964) erectedHystrichosphaera bentorii var truncata to encompass speci-mens with shorter processes This variety later was trans-ferred along with the species to Spiniferites (Lentin ampWilliams 1973) Spiniferites bentorii var globus Morzadec-Kerfourn 1979 p 222ndash224 pl 31 fig 10 was introduced toencompass specimens with a spheroidal central body asmaller apical boss and longer more slender processesSeveral subspecies have been described from the Neogenedeposits of the Pannonian Basin in central Europe of whichfour were validly published Spiniferites bentorii subsp buda-jenoensis Seurouto-Szentai 1986 Spiniferites bentorii subsp granu-latus Fuchs amp Seurouto-Szentai 1991 Spiniferites bentorii subspoblongus Seurouto-Szentai 1986 (now Spiniferites oblongusSoliman amp Riding 2017) and Spiniferites bentorii subsp pan-nonicus Seurouto-Szentai 1986 Another four were proposed butnot validly published ldquoSpiniferites bentorii subsp coniunctusrdquoSeurouto-Szentai 1990 ldquoSpiniferites bentorii subsp matraensisrdquo Seurouto-Szentai 1988 ldquoSpiniferites bentorii subsp piriformisrdquo Seurouto-Szentai1988 and ldquoSpiniferites bentorii subsp pseudooblongusrdquo Seurouto-Szentai 1983 None of the eight morphotypes have beenrecorded from PliocenendashQuaternary sedimentsComparison Spiniferites bentorii is similar to Spiniferites lazusin that it can have fenestrate process bases but is distin-guished from Spiniferites lazus by its larger size pear-shape(as opposed to elongate-ovoid) ambitus process form andcingulum displacement (Reid 1974) According to Rochonet al (1999) the central body of Spiniferites bentorii can beovoid but herein we assign such ovoid forms to Spiniferiteslazus Spiniferites bentorii differs from Spiniferites multisphae-rus on the basis of its wall structure which does not containvacuoles unlike Spiniferites multisphaerus (Price amp Pospelova2014) Spiniferites hainanensis has an ovoid central body ForSpiniferites nodosus see Section 226Remarks During the first workshop VP pointed out thatSpiniferites bentorii can have intergonal processes specimensillustrated by Price amp Pospelova (2014) corresponding toSpiniferites bentorii subsp truncatus show such processesRochon et al (1999) also mentioned the presence of occa-sional intergonal processes in this species LL remarked thatthe presence of intergonal processes is not a problem at thespecies level if they are rare or sparse and there is no morethan one between adjacent gonal processes LL furtherremarked that he does not consider the apical boss animportant characteristic but that the pear-shaped centralbody is important Wall amp Dale (1970) while transferring

18 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Distinguishing characters Ovoid central body with a pro-nounced apical boss and lacking sutural ornamentation Theouter surface is smooth and bears 24 lobate branched exclu-sively gonal processes which can have fenestrate bases Theprocesses are hollow or sometimes partly solid Their distalends expand into lobate short branches and some have irregu-larly fringed tips Two of the apical processes are fused andhave a flattened distal end which can also be the case for twoof the antapical processes Expressed tabulation is 40 600 6cs5ndash6000 1p 100 00 The ends of the cingulum are displaced byapparently about one cingulum width and the sulcus isstraight The archeopyle is formed by loss of plate 3rdquo and theoperculum is free (Based on Demetrescu 1989 p 51ndash53)Dimensions Central body length 47ndash50 mm width 32ndash35mmthickness 33ndash35 mm apical boss 1ndash7 mm long process length15ndash26 mm (Demetrescu 1989)Biological affinity UnknownIntraspecific morphotypes NoneComparison Achomosphaera argesensis differs fromAchomosphaera ramulifera in having an apical boss whichresembles that of Achomosphaera improcera Islam 1983 fromthe Eocene and Spiniferites bentorii but Achomosphaeraimprocera has much shorter processes and Spiniferites ben-torii has low sutures Achomosphaera argesensis also resem-bles Achomosphaera andalousiensis but the latter does nothave a flat apical process or processes with lobate tipsSpiniferites validus Seurouto-Szentai 1982 described from thePannonian Basin differs from Achomosphaera argesensis inhaving a thin elongated apical process instead of a largeflattened apical process (Based on Demetrescu 1989)Remarks PJM was unsuccessful in locating the holotype inBucharest because the Institute of Geology Bucharest hasbeen dismantled and the location of the type slides is nolonger known moreover it was not possible to contactEmanuel Demetrescu so the holotype is effectively lostEaton (1996 pl IV figs 6ndash7) illustrated well-preserved speci-mens from the Black Sea with membranes connecting thedistal ends of apical and antapical gonal processes Eatonrsquosslides are archived in the Palynological Slide Collection ofthe Department of Paleontology at the Natural HistoryMuseum London UK The age of Eatonrsquos sample is uncer-tain VT noted that he saw specimens of Achomosphaeraargesensis from the Pannonian Basin However based on theoriginal publication of Demetrescu (1989) VT identifiedAchomosphaera argesensis on the basis that the holotypeillustrations show clear membranes connecting the gonalprocesses associated with the apical plates He suggestedthat this is the criterion to consider in separatingAchomosphaera argesensis from the other species describedfrom the Pannonian BasinParatethys AG agreed that it isvery distinct and suggested that these forms ofAchomosphaera may fit better in Spiniferites During the draft-ing of this report LL remarked that a lack of true septabetween most processes would still leave Achomosphaeraargesensis in Achomosphaera FS noted during drafting thatshe observed Achomosphaera argesensis in Upper Miocene(regional Pontian Stage) sediments of the ParatethyanBlack Sea

23 Achomosphaera callosa Matsuoka 1983 (Plate 2figures 1ndash3)

Synonymy NoneHolotype Matsuoka 1983 pl 11 figs 6andashcType locality Ota Tsugawa-cho Niigata Prefecture cen-tral JapanType stratum Tokonami Formation equivalent to theNishiyama Formation Pliocene (Matsuoka 1983)Etymology From Latin callosa thick skinned in reference tothe thick wall (Matsuoka 1983)Distinguishing characters Central body thick-walled spher-oidal to (rarely) ovoid with a coarsely granular outer surfaceThe smooth processes are gonal only and sutural septa areonly occasionally present The archeopyle is formed by lossof plate 3rdquo and is reduced (Based on Matsuoka 1983p 128ndash129)Dimensions Central body length 36ndash53 mm width 36ndash45 mmwall thickness up to 3 mm process length up to 15 mm(Matsuoka 1983 p 128)Biological affinity UnknownIntraspecific morphotypes NoneComparison Achomosphaera callosa resemblesAchomosphaera crassipellis (Deflandre amp Cookson 1955)Stover amp Evitt 1978 and Achomosphaera cf sagena Davey ampWilliams 1966 of Gocht (1969 p 36 pl 7 figs 1ndash2) but dif-fers from these two forms in having a coarsely granular outersurface and exclusively gonal processes (Matsuoka 1983) Thecentral body of Achomosphaera crassipellis is larger(74ndash89 mm) than that of Achomosphaera callosa and has athicker wall judging from the illustration in Deflandre ampCookson (1955 no measurements provided) and longer proc-esses (23ndash26 mm) (measurements from Deflandre ampCookson 1955)Remarks Matsuoka (1983 p 128ndash129) describedAchomosphaera callosa from the Pliocene and LowerPleistocene of Japan LL expressed that he has no problemwith the species although he has not observed it inQuaternary sediments The exact range of the speciesremains to be determined MJH remarked that the centralbody apparently has a solid pedium a prismatic layer on topand open luxuria but no tegillum This leads to the questionof what the processes are made of an issue probably bestanswered through scanning electron microscopy Other thanthat MJH considers Achomosphaera callosa a distinctive spe-cies and LL agrees

24 Achomosphaera granulata Mao Shaozhi 1989

Synonymy Non Achomosphaera sp A of Matsuoka 1983 pl11 figs 1ndash5 illustrated in Plate 2 figures 4ndash8Holotype Mao Shaozhi 1989 pl 28 fig 10Type locality Northern Xisha Trench South China SeaType stratum Upper Pleistocene (Mao Shaozhi 1989)Etymology Derived from the occurrence of dense granuleson the wall and processes (Mao Shaozhi 1989)Distinguishing characters Central body ovoid thick-walledlight brown to brown commonly with a short apical boss

PALYNOLOGY 13

The apical horn has a basal width greater than its length(typically 3ndash5 mm) and has a truncated distal end The cingu-lum is 5ndash7mm wide delimited by dense granules separatingthe cyst into a sub-triangular epicyst and rounded to nearlytrapezoidal hypocyst There are two wall layers the thick

outer wall forming the exclusively gonal processes The sur-face of the outer wall is ornamented by dense and well-dis-tributed granules The granules are coarse and sometimeslink to form curved lines They are developed on both thecentral body and processes and make the outline of the

Plate 2 1ndash3 Holotype of Achomosphaera callosa Matsuoka 1983 at high to low focus 4 Topotype 5 Focus on holotype showing detail of wall texture and pro-cess 6ndash8 Upper to lower focus of Achomosphaera sp A of Matsuoka (1983) plate 11 figures 1ndash5 [Mao Shaozhi (1989) considered her specimens ofAchomosphaera granulata the same as these specimens from the Lower to Upper Miocene of the Niigata district (central Japan) but we disagree with this assess-ment] Slides provided by KM photographed by KNM All scale bars frac14 10mm

14 K N MERTENS ET AL

processes rough The processes can be perforated due to cor-rosion according to Mao Shaozhi (1989) The processes havewide bases and taper rapidly No sutures are present exceptfor the granules that mark the cingulum dorsally The archeo-pyle is formed by loss of plate 3rdquo (Based on Mao Shaozhi1989 p 139 translated from Chinese by HG and observationsof Achomosphaera sp A of Matsuoka 1983 p 130)Dimensions Central body length (including apical horn)45ndash53 mm width 37ndash45 mm wall thickness 2 mm processlength 10ndash13 mm (Mao Shaozhi 1989)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species can be distinguished from all simi-lar Spiniferites and Achomosphaera species by the uniquedense granules covering the wall and processes its thickouter wall and brown colour (Mao Shaozhi 1989)Remarks Mao Shaozhi (1989) considered Achomosphaeragranulata the same as Achomosphaera sp A of Matsuoka1983 from the Lower to Upper Miocene of the Niigata dis-trict central Japan but we disagree with this assessmentsee also Londeix et al (2018)

The species has been recorded only in Quaternary sedi-ments of the South China Sea by Mao Shaozhi (1989) andMao Shaozhi amp Harland (1993) Londeix et al (2018) consid-ered this species to belong to Hafniasphaera others includ-ing KNM and VP were not convinced because the publishedholotype images are not sharp and the description does notmention the presence of vacuoles

25 Achomosphaera ramosasimilis (Yun Hyesu 1981)Londeix et al 1999

Synonymy Achomosphaera ramulifera subsp ramosasimilisYun Hyesu 1981 p 14ndash15 pl 1 figs 1 8 text-fig 3b

Spiniferites ramuliferus (auct non Deflandre) Reid Reid1974 p 608 pl 4 figs 39ndash40Holotype Yun Hyesu 1981 pl 1 fig 1 text-fig 3b Fensomeet al 1991 figs 1ndash2 p 719 fig 4 p 719 amp 721 [initiallydescribed in German by Yun Hyesu 1981 translated byFensome et al 1991]Type locality Timmermann brickyard new pit near EsbeckWestphalia GermanyType stratum Santonian Upper Cretaceous (Yun Hyesu 1981)Etymology In reference to the close similarity between thisspecies and Spiniferites ramosus (Fensome et al 1991 p 721)Distinguishing characters Thick-walled ovoid to spheroidalcentral body with a granular or smooth pedium and asmooth tegillum bearing hollow distally closed exclusivelygonal processes The processes on the cingulum may be con-nected by crests similar to some processes in the apical andantapical areas There is always a thin apical process occa-sionally with a terminal hooklet Sutures can be partly pre-sent The archeopyle is formed by the loss of plate 3rdquo (Basedon Yun Hyesu 1981 p 14ndash15) and Fensome et al 1991p 719ndash720)Dimensions Central body length 30 (32) 36 mm width 32(39) 42 mm maximum length of processes 16ndash18 mm (YunHyesu 1981)

Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Yun Hyesu (1981) becauseAchomosphaera ramosasimilis possesses a thick wall partialpresence of sutures does not have all processes connectedby crests bears a simple apical process and no antapicalbranched process it differs from Achomosphaera ramuliferaRemarks This species was observed by Londeix et al (1999)from the Lower Pliocene of the Strait of Sicily centralMediterranean Sea LL also considers specimens ofldquoSpiniferites ramuliferusrdquo described by Reid (1974) from recentraised-beach deposits at Woodgrange Northern Ireland UKto be probably Achomosphaera ramosasimilis and notAchomosphaera ramulifera see discussion in Londeix et al(2018) and Gurdebeke et al (2018)

26 Spiniferites alaskensis Marret et al 2001 ex Marretin Fensome amp Williams 2004

Synonymy NoneHolotype Marret et al 2001 pl 1 figs 7ndash9 designated byMarret in Fensome amp Williams (2004)Type locality ODP Site 887 Gulf of Alaska northeasternNorth Pacific Gulf of AlaskaType stratum Marine Isotope Substage 5e Sample 887B-2H-5 65 cm (Marret et al 2001)Etymology Named alaskensis from the type locality (Marretet al 2001)Distinguishing characters Ovoid shape with an apical bossThe cyst wall is thin (lt 1 mm) and has a finely granulate toscabrate surface Processes are gonal and relatively broad(2 mm) each process terminating in a simple trifurcation withpointed ends Low sutural septa are present between proc-esses and define a gonyaulacacean tabulation The archeo-pyle is formed by loss of plate 3rdquo (Based on Marret et al2001 p 384ndash385)Dimensions Central body length 263 (314) 368 mm width236 (293) 315 mm process length 75 (101) 125 mm (Marretet al 2001)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species differs from other Spiniferites spe-cies only by the process shape and termination Spiniferitesalaskensis has broad processes with simple short trifurcatebranches with pointed ends such processes are unusual forlate Quaternary Spiniferites species (Marret et al 2001p 386)Remarks The name was validated by Marret in Fensome ampWilliams (2004) because Marret et al (2001) did not indicatewhich of the illustrations represented the holotype This spe-cies is discussed in Marret amp Mertens (2018)

27 Spiniferites asperulus Matsuoka 1983 (Plate 3figures 1ndash6)

Synonymy NoneHolotype Matsuoka 1983 pl 12 fig 2

PALYNOLOGY 15

Plate 3 1ndash3 Holotype of Spiniferites asperulus Matsuoka 1983 in ventral view at high to low focus 4 Same specimen mid-focus showing wall structure 5 Samespecimen mid-focus showing antapical processes 6ndash8 Holotype of Spiniferites firmus Matsuoka 1983 in ventral view at high to low focus 9 Same specimen show-ing microgranular wall 10 Same specimen showing distal ends of cingular processes Slides provided by KM photographed by KNM All scale bars frac14 10 mmexcept 1ndash3 7ndash9frac14 20 mm

16 K N MERTENS ET AL

Type locality Nishiyama Nishiyama-cho Niigata PrefectureCentral JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin asper (somewhat rough) in ref-erence to the coarsely granular surface of the cyst (Matsuoka1983 p 131)Distinguishing characters The subspheroidal to ovoid cysthas a relatively thick wall The tegillum forms many smallinvaginations in the intratabular area No granules are pre-sent The processes are slender membranous gonal (andoccasionally intergonal) and widen at the base the twoantapical processes are somewhat more robust and con-nected by a slightly elevated crest (Based on Matsuoka1983 p 131ndash132 and observations by KNM)Dimensions Central body length 48ndash69 mm width45ndash64 mm process length up to 16 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison The wall structure is similar to that ofSpiniferites ludhamensis but there are many more invagina-tions in Spiniferites asperulus and Spiniferites ludhamensis issmaller (central body length 38ndash49 mm) and has completelyhollow processes Spiniferites membranaceus has a granu-lar wallRemarks LL expressed the difficulty in differentiating thisspecies from others such as Spiniferites firmus KM stated thatafter having described it back in 1983 he now has no clearrecollection of what this species is LL looked at the holotypebut adds that he did not look at the total population andtherefore finds it hard to say if this is a distinct species or notHe noted that granulation which may occur as a variation inmany species could be a response to specific environmentalconditions The original description mentions the occasionaloccurrence of intergonal processes a feature confirmed byKM This species is discussed in Limoges et al (2018)

28 Spiniferites belerius Reid 1974

Synonymy Non Spiniferites belerius sensu Harland 1977 p98ndash99 pl 1 figs 7ndash10 pl 2 figs 7ndash10 16ndash21 25ndash57 [frac14Spiniferites membranaceus]Holotype Reid 1974 pl 2 figs 12ndash13Type locality Severn Estuary England UKType stratum Modern surface sediments (Reid 1974)Etymology From the Latin belerium which is the Romanname for the promontory on which Landrsquos End CornwallEngland is situated (Reid 1974 p 596)Distinguishing characters The small central body is ovalwith a finely granular wall and an apical boss The processes aregonal and connected by low crests with a typical process shapeThis species was described as having a characteristic largelsquotrumpet-shapedrsquo posterior process at the junction of plates 1and 2 (Reid 1974 p 597 but see Gurdebeke et al 2018)Dimensions Central body length 35ndash42 mm width28ndash37 mm maximum process length 7ndash10 mm posterior pro-cess length 10ndash15 mm (Reid 1974)

Biological affinity Gonyaulax scrippsae Kofoid 1911 asdepicted by Wall amp Dale (1968 p 270 pl 1 fig 14) accord-ing to Reid (1974) LL and KZ noted some similarity to thecyst of Gonyaulax baltica Ellegaard Lewis amp Harding 2002 asproduced in batch-cultures and illustrated by Ellegaard et al(2002) KZ felt she could not understand the cyst-based spe-cies concept here because of the large morphological vari-ation documented in that study VP wondered if anyonecould distinguish cysts of Gonyaulax baltica in the sedimentsME replied that some cysts of Gonyaulax baltica could be thesame as Spiniferites belerius but ideally we should first deriveSpiniferites belerius in culture and compare the phylogeneticsIntraspecific morphotypes NoneComparison LL noted during the first workshop thatSpiniferites bentorii is larger overall and has a hypocyst largerthan the epicyst Spiniferites belerius is similar to Spiniferitesconiconcavus but the latter has wider process basesRemarks During the first and second workshops there wasagreement that this species is problematic LL consideredthis species as having a large morphological variation Healso suggested that this species is defined by central bodysize and shape and the presence of short processes that arenot well developed in the apical region PJM mentioned thatspecimens from the Black Sea show a very wide morpho-logical range and that these could be used as an illustrationof the speciesrsquo various morphologies KNM stated howeverthat potentially different species have been described asSpiniferites belerius given the great variation that he hasobserved globally PJM remarked that this is hard to tellwithout culture studies LL proposed that two categories bedefined for practical purposes typical and atypical morpho-types These categories can then be used in paleoecologicalstudies LL further underlined the wide range in morphologythat appears to characterise many Paratethyan species PJMasked LL whether he considered Spiniferites belerius to beParatethyan and LL answered ldquonordquo He noted that there is aclear regular shape and a wide range of morphological vari-ability and that it is sometimes hard to know if this repre-sents infraspecific variability or more than one species WhenPJM asked what LL considers a Paratethyan species hereplied that it is an ldquoendemicrdquo species with a clearly definedtypical morphology He used Seriliodinium explicatum Eaton1996 Invertocysta sp A of Londeix et al (2009)Invertocysta sp B of Londeix et al (2009) Pterocysta crucifor-mis Rochon et al 2003 Pyxidinopsis psilata (Wall amp Dale inWall et al 1973) Head 1994 Galeacysta etrusca Corradini ampBiffi 1988 and Caspidinium rugosum Marret in Marret et al2004 as good examples of Paratethyan species PJM com-mented in draft that LLrsquos examples include taxa not recordedoutside the Ponto-Caspian basins together with taxa havingwide ranges from the saline western Mediterranean Sea tothe brackish Aral Sea She considers it difficult to justify theword endemic for such a broad grouping of taxa She con-siders that by definition an endemic species means native orrestricted to a certain country or area and should not beused for Paratethyan forms as the Paratethys was a vastEuro-Asian waterway BD suggested that we should decidewhether we can put species into two different groups those

PALYNOLOGY 17

that are stable and well-described and those that originatefrom particular extreme environments that induce increasedecological pressure He said that much of the discussionappears to be focused on forms from extreme environmentsHe noted however that he does not know how to applysuch an approach in practice but proposed that ldquoweirdrdquomorphotypes should be addressed ldquolater onrdquo once clearbasic forms have been established KNM suggested that thetopotype material of Spiniferites belerius should be looked atto evaluate the morphological variability present This hasnow been done by Gurdebeke et al (2018) Limoges et al(2018) have accepted the presence of additional intergonalprocesses in this species

29 Spiniferites bentorii (Rossignol 1964) Wall ampDale 1970

Synonymy Hystrichosphaera bentorii Rossignol 1964 p84ndash85 pl 1 figs 3 3 inset 5ndash8 pl 3 figs 2ndash3 text-figs AndashF

Leptodinium churchillii Harland 1968 p 548 550ndash551 figs12ndash13 22ndash24

non Spiniferites nodosus (Wall 1967) Sarjeant 1970[Synonymy proposed by Reid (1974 p 599) is here rejected]

non Spiniferites bentorii sensu Wall amp Dale 1970 p 52 pl1 figs 26 28Holotype Rossignol 1964 pl 1 figs 3 7ndash8Type locality Ashdod borehole coastal plain IsraelType stratum Quaternary sediments 172ndash1725m depth(Rossignol 1964)Etymology Named after Dr Yaakov Ben-Tor (b 1910 ndash d2002) Israeli geologist who was head of the IsraeliGeological Survey at the time of Martine Rossignolrsquos studyand provided her with samplesDistinguishing characters This species typically has a pear-shaped central body with a characteristic apical boss It bearscharacteristic tapering slender gonal and occasionally inter-gonal processes with the two antapical processes being thelongest The process bases may be fenestrate Sutures aremarked by low ridges and sometimes vacuoles are presentin the sutures The wall is smooth to microgranular The cin-gular displacement is relatively large (three times its widthaccording to Reid 1974 one and a half to two times itswidth according to Wall 1965) Tabulation is typical for thegenus according to Wall (1965) and Harland (1968) with fourapical plates although the suture between 1 and 2 is faintand was not observed by Rossignol (1964) or Price ampPospelova (2014) both interpreting this as indicating thepresence of only three apical plates visible on the cyst Thesulcus is often well expressed and straight and widens poster-iorly (Reid 1974) The archeopyle is formed by loss of plate 3rdquoand is reduced with rounded corners (Based on Rossignol1964 p 84ndash85 Harland 1968 p 542ndash543 Reid 1974p 598ndash600 Rochon et al 1999 p 34 Price amp Pospelova2014 p 13 and observations of the participants) MJH notedin draft that specimens from the Last Interglacial of theBaltic have distinctive lateral cingular processes in which thedistal furcations branch abruptly at 90 to the process shafts

and point towards both poles (Head 2007 fig 7jkm)MJH further noted in draft that specimens of S bentorii thathe has observed have archeopyles well defined bythe suturesDimensions Central body length 60ndash73 mm width45ndash63 mm process length 15ndash20 mm (25 mm for the antapi-cal processes) (Rossignol 1964) Central body length58ndash69 mm width 52ndash55 mm length of processes 0ndash20 mm(Reid 1974)Biological affinity Gonyaulax digitale (Pouchet 1883) Kofoid1911 according to Wall amp Dale (1967 p 352) and Dodge(1989 p 283)Intraspecific morphotypes Rossignol (1964) erectedHystrichosphaera bentorii var truncata to encompass speci-mens with shorter processes This variety later was trans-ferred along with the species to Spiniferites (Lentin ampWilliams 1973) Spiniferites bentorii var globus Morzadec-Kerfourn 1979 p 222ndash224 pl 31 fig 10 was introduced toencompass specimens with a spheroidal central body asmaller apical boss and longer more slender processesSeveral subspecies have been described from the Neogenedeposits of the Pannonian Basin in central Europe of whichfour were validly published Spiniferites bentorii subsp buda-jenoensis Seurouto-Szentai 1986 Spiniferites bentorii subsp granu-latus Fuchs amp Seurouto-Szentai 1991 Spiniferites bentorii subspoblongus Seurouto-Szentai 1986 (now Spiniferites oblongusSoliman amp Riding 2017) and Spiniferites bentorii subsp pan-nonicus Seurouto-Szentai 1986 Another four were proposed butnot validly published ldquoSpiniferites bentorii subsp coniunctusrdquoSeurouto-Szentai 1990 ldquoSpiniferites bentorii subsp matraensisrdquo Seurouto-Szentai 1988 ldquoSpiniferites bentorii subsp piriformisrdquo Seurouto-Szentai1988 and ldquoSpiniferites bentorii subsp pseudooblongusrdquo Seurouto-Szentai 1983 None of the eight morphotypes have beenrecorded from PliocenendashQuaternary sedimentsComparison Spiniferites bentorii is similar to Spiniferites lazusin that it can have fenestrate process bases but is distin-guished from Spiniferites lazus by its larger size pear-shape(as opposed to elongate-ovoid) ambitus process form andcingulum displacement (Reid 1974) According to Rochonet al (1999) the central body of Spiniferites bentorii can beovoid but herein we assign such ovoid forms to Spiniferiteslazus Spiniferites bentorii differs from Spiniferites multisphae-rus on the basis of its wall structure which does not containvacuoles unlike Spiniferites multisphaerus (Price amp Pospelova2014) Spiniferites hainanensis has an ovoid central body ForSpiniferites nodosus see Section 226Remarks During the first workshop VP pointed out thatSpiniferites bentorii can have intergonal processes specimensillustrated by Price amp Pospelova (2014) corresponding toSpiniferites bentorii subsp truncatus show such processesRochon et al (1999) also mentioned the presence of occa-sional intergonal processes in this species LL remarked thatthe presence of intergonal processes is not a problem at thespecies level if they are rare or sparse and there is no morethan one between adjacent gonal processes LL furtherremarked that he does not consider the apical boss animportant characteristic but that the pear-shaped centralbody is important Wall amp Dale (1970) while transferring

18 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

The apical horn has a basal width greater than its length(typically 3ndash5 mm) and has a truncated distal end The cingu-lum is 5ndash7mm wide delimited by dense granules separatingthe cyst into a sub-triangular epicyst and rounded to nearlytrapezoidal hypocyst There are two wall layers the thick

outer wall forming the exclusively gonal processes The sur-face of the outer wall is ornamented by dense and well-dis-tributed granules The granules are coarse and sometimeslink to form curved lines They are developed on both thecentral body and processes and make the outline of the

Plate 2 1ndash3 Holotype of Achomosphaera callosa Matsuoka 1983 at high to low focus 4 Topotype 5 Focus on holotype showing detail of wall texture and pro-cess 6ndash8 Upper to lower focus of Achomosphaera sp A of Matsuoka (1983) plate 11 figures 1ndash5 [Mao Shaozhi (1989) considered her specimens ofAchomosphaera granulata the same as these specimens from the Lower to Upper Miocene of the Niigata district (central Japan) but we disagree with this assess-ment] Slides provided by KM photographed by KNM All scale bars frac14 10mm

14 K N MERTENS ET AL

processes rough The processes can be perforated due to cor-rosion according to Mao Shaozhi (1989) The processes havewide bases and taper rapidly No sutures are present exceptfor the granules that mark the cingulum dorsally The archeo-pyle is formed by loss of plate 3rdquo (Based on Mao Shaozhi1989 p 139 translated from Chinese by HG and observationsof Achomosphaera sp A of Matsuoka 1983 p 130)Dimensions Central body length (including apical horn)45ndash53 mm width 37ndash45 mm wall thickness 2 mm processlength 10ndash13 mm (Mao Shaozhi 1989)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species can be distinguished from all simi-lar Spiniferites and Achomosphaera species by the uniquedense granules covering the wall and processes its thickouter wall and brown colour (Mao Shaozhi 1989)Remarks Mao Shaozhi (1989) considered Achomosphaeragranulata the same as Achomosphaera sp A of Matsuoka1983 from the Lower to Upper Miocene of the Niigata dis-trict central Japan but we disagree with this assessmentsee also Londeix et al (2018)

The species has been recorded only in Quaternary sedi-ments of the South China Sea by Mao Shaozhi (1989) andMao Shaozhi amp Harland (1993) Londeix et al (2018) consid-ered this species to belong to Hafniasphaera others includ-ing KNM and VP were not convinced because the publishedholotype images are not sharp and the description does notmention the presence of vacuoles

25 Achomosphaera ramosasimilis (Yun Hyesu 1981)Londeix et al 1999

Synonymy Achomosphaera ramulifera subsp ramosasimilisYun Hyesu 1981 p 14ndash15 pl 1 figs 1 8 text-fig 3b

Spiniferites ramuliferus (auct non Deflandre) Reid Reid1974 p 608 pl 4 figs 39ndash40Holotype Yun Hyesu 1981 pl 1 fig 1 text-fig 3b Fensomeet al 1991 figs 1ndash2 p 719 fig 4 p 719 amp 721 [initiallydescribed in German by Yun Hyesu 1981 translated byFensome et al 1991]Type locality Timmermann brickyard new pit near EsbeckWestphalia GermanyType stratum Santonian Upper Cretaceous (Yun Hyesu 1981)Etymology In reference to the close similarity between thisspecies and Spiniferites ramosus (Fensome et al 1991 p 721)Distinguishing characters Thick-walled ovoid to spheroidalcentral body with a granular or smooth pedium and asmooth tegillum bearing hollow distally closed exclusivelygonal processes The processes on the cingulum may be con-nected by crests similar to some processes in the apical andantapical areas There is always a thin apical process occa-sionally with a terminal hooklet Sutures can be partly pre-sent The archeopyle is formed by the loss of plate 3rdquo (Basedon Yun Hyesu 1981 p 14ndash15) and Fensome et al 1991p 719ndash720)Dimensions Central body length 30 (32) 36 mm width 32(39) 42 mm maximum length of processes 16ndash18 mm (YunHyesu 1981)

Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Yun Hyesu (1981) becauseAchomosphaera ramosasimilis possesses a thick wall partialpresence of sutures does not have all processes connectedby crests bears a simple apical process and no antapicalbranched process it differs from Achomosphaera ramuliferaRemarks This species was observed by Londeix et al (1999)from the Lower Pliocene of the Strait of Sicily centralMediterranean Sea LL also considers specimens ofldquoSpiniferites ramuliferusrdquo described by Reid (1974) from recentraised-beach deposits at Woodgrange Northern Ireland UKto be probably Achomosphaera ramosasimilis and notAchomosphaera ramulifera see discussion in Londeix et al(2018) and Gurdebeke et al (2018)

26 Spiniferites alaskensis Marret et al 2001 ex Marretin Fensome amp Williams 2004

Synonymy NoneHolotype Marret et al 2001 pl 1 figs 7ndash9 designated byMarret in Fensome amp Williams (2004)Type locality ODP Site 887 Gulf of Alaska northeasternNorth Pacific Gulf of AlaskaType stratum Marine Isotope Substage 5e Sample 887B-2H-5 65 cm (Marret et al 2001)Etymology Named alaskensis from the type locality (Marretet al 2001)Distinguishing characters Ovoid shape with an apical bossThe cyst wall is thin (lt 1 mm) and has a finely granulate toscabrate surface Processes are gonal and relatively broad(2 mm) each process terminating in a simple trifurcation withpointed ends Low sutural septa are present between proc-esses and define a gonyaulacacean tabulation The archeo-pyle is formed by loss of plate 3rdquo (Based on Marret et al2001 p 384ndash385)Dimensions Central body length 263 (314) 368 mm width236 (293) 315 mm process length 75 (101) 125 mm (Marretet al 2001)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species differs from other Spiniferites spe-cies only by the process shape and termination Spiniferitesalaskensis has broad processes with simple short trifurcatebranches with pointed ends such processes are unusual forlate Quaternary Spiniferites species (Marret et al 2001p 386)Remarks The name was validated by Marret in Fensome ampWilliams (2004) because Marret et al (2001) did not indicatewhich of the illustrations represented the holotype This spe-cies is discussed in Marret amp Mertens (2018)

27 Spiniferites asperulus Matsuoka 1983 (Plate 3figures 1ndash6)

Synonymy NoneHolotype Matsuoka 1983 pl 12 fig 2

PALYNOLOGY 15

Plate 3 1ndash3 Holotype of Spiniferites asperulus Matsuoka 1983 in ventral view at high to low focus 4 Same specimen mid-focus showing wall structure 5 Samespecimen mid-focus showing antapical processes 6ndash8 Holotype of Spiniferites firmus Matsuoka 1983 in ventral view at high to low focus 9 Same specimen show-ing microgranular wall 10 Same specimen showing distal ends of cingular processes Slides provided by KM photographed by KNM All scale bars frac14 10 mmexcept 1ndash3 7ndash9frac14 20 mm

16 K N MERTENS ET AL

Type locality Nishiyama Nishiyama-cho Niigata PrefectureCentral JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin asper (somewhat rough) in ref-erence to the coarsely granular surface of the cyst (Matsuoka1983 p 131)Distinguishing characters The subspheroidal to ovoid cysthas a relatively thick wall The tegillum forms many smallinvaginations in the intratabular area No granules are pre-sent The processes are slender membranous gonal (andoccasionally intergonal) and widen at the base the twoantapical processes are somewhat more robust and con-nected by a slightly elevated crest (Based on Matsuoka1983 p 131ndash132 and observations by KNM)Dimensions Central body length 48ndash69 mm width45ndash64 mm process length up to 16 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison The wall structure is similar to that ofSpiniferites ludhamensis but there are many more invagina-tions in Spiniferites asperulus and Spiniferites ludhamensis issmaller (central body length 38ndash49 mm) and has completelyhollow processes Spiniferites membranaceus has a granu-lar wallRemarks LL expressed the difficulty in differentiating thisspecies from others such as Spiniferites firmus KM stated thatafter having described it back in 1983 he now has no clearrecollection of what this species is LL looked at the holotypebut adds that he did not look at the total population andtherefore finds it hard to say if this is a distinct species or notHe noted that granulation which may occur as a variation inmany species could be a response to specific environmentalconditions The original description mentions the occasionaloccurrence of intergonal processes a feature confirmed byKM This species is discussed in Limoges et al (2018)

28 Spiniferites belerius Reid 1974

Synonymy Non Spiniferites belerius sensu Harland 1977 p98ndash99 pl 1 figs 7ndash10 pl 2 figs 7ndash10 16ndash21 25ndash57 [frac14Spiniferites membranaceus]Holotype Reid 1974 pl 2 figs 12ndash13Type locality Severn Estuary England UKType stratum Modern surface sediments (Reid 1974)Etymology From the Latin belerium which is the Romanname for the promontory on which Landrsquos End CornwallEngland is situated (Reid 1974 p 596)Distinguishing characters The small central body is ovalwith a finely granular wall and an apical boss The processes aregonal and connected by low crests with a typical process shapeThis species was described as having a characteristic largelsquotrumpet-shapedrsquo posterior process at the junction of plates 1and 2 (Reid 1974 p 597 but see Gurdebeke et al 2018)Dimensions Central body length 35ndash42 mm width28ndash37 mm maximum process length 7ndash10 mm posterior pro-cess length 10ndash15 mm (Reid 1974)

Biological affinity Gonyaulax scrippsae Kofoid 1911 asdepicted by Wall amp Dale (1968 p 270 pl 1 fig 14) accord-ing to Reid (1974) LL and KZ noted some similarity to thecyst of Gonyaulax baltica Ellegaard Lewis amp Harding 2002 asproduced in batch-cultures and illustrated by Ellegaard et al(2002) KZ felt she could not understand the cyst-based spe-cies concept here because of the large morphological vari-ation documented in that study VP wondered if anyonecould distinguish cysts of Gonyaulax baltica in the sedimentsME replied that some cysts of Gonyaulax baltica could be thesame as Spiniferites belerius but ideally we should first deriveSpiniferites belerius in culture and compare the phylogeneticsIntraspecific morphotypes NoneComparison LL noted during the first workshop thatSpiniferites bentorii is larger overall and has a hypocyst largerthan the epicyst Spiniferites belerius is similar to Spiniferitesconiconcavus but the latter has wider process basesRemarks During the first and second workshops there wasagreement that this species is problematic LL consideredthis species as having a large morphological variation Healso suggested that this species is defined by central bodysize and shape and the presence of short processes that arenot well developed in the apical region PJM mentioned thatspecimens from the Black Sea show a very wide morpho-logical range and that these could be used as an illustrationof the speciesrsquo various morphologies KNM stated howeverthat potentially different species have been described asSpiniferites belerius given the great variation that he hasobserved globally PJM remarked that this is hard to tellwithout culture studies LL proposed that two categories bedefined for practical purposes typical and atypical morpho-types These categories can then be used in paleoecologicalstudies LL further underlined the wide range in morphologythat appears to characterise many Paratethyan species PJMasked LL whether he considered Spiniferites belerius to beParatethyan and LL answered ldquonordquo He noted that there is aclear regular shape and a wide range of morphological vari-ability and that it is sometimes hard to know if this repre-sents infraspecific variability or more than one species WhenPJM asked what LL considers a Paratethyan species hereplied that it is an ldquoendemicrdquo species with a clearly definedtypical morphology He used Seriliodinium explicatum Eaton1996 Invertocysta sp A of Londeix et al (2009)Invertocysta sp B of Londeix et al (2009) Pterocysta crucifor-mis Rochon et al 2003 Pyxidinopsis psilata (Wall amp Dale inWall et al 1973) Head 1994 Galeacysta etrusca Corradini ampBiffi 1988 and Caspidinium rugosum Marret in Marret et al2004 as good examples of Paratethyan species PJM com-mented in draft that LLrsquos examples include taxa not recordedoutside the Ponto-Caspian basins together with taxa havingwide ranges from the saline western Mediterranean Sea tothe brackish Aral Sea She considers it difficult to justify theword endemic for such a broad grouping of taxa She con-siders that by definition an endemic species means native orrestricted to a certain country or area and should not beused for Paratethyan forms as the Paratethys was a vastEuro-Asian waterway BD suggested that we should decidewhether we can put species into two different groups those

PALYNOLOGY 17

that are stable and well-described and those that originatefrom particular extreme environments that induce increasedecological pressure He said that much of the discussionappears to be focused on forms from extreme environmentsHe noted however that he does not know how to applysuch an approach in practice but proposed that ldquoweirdrdquomorphotypes should be addressed ldquolater onrdquo once clearbasic forms have been established KNM suggested that thetopotype material of Spiniferites belerius should be looked atto evaluate the morphological variability present This hasnow been done by Gurdebeke et al (2018) Limoges et al(2018) have accepted the presence of additional intergonalprocesses in this species

29 Spiniferites bentorii (Rossignol 1964) Wall ampDale 1970

Synonymy Hystrichosphaera bentorii Rossignol 1964 p84ndash85 pl 1 figs 3 3 inset 5ndash8 pl 3 figs 2ndash3 text-figs AndashF

Leptodinium churchillii Harland 1968 p 548 550ndash551 figs12ndash13 22ndash24

non Spiniferites nodosus (Wall 1967) Sarjeant 1970[Synonymy proposed by Reid (1974 p 599) is here rejected]

non Spiniferites bentorii sensu Wall amp Dale 1970 p 52 pl1 figs 26 28Holotype Rossignol 1964 pl 1 figs 3 7ndash8Type locality Ashdod borehole coastal plain IsraelType stratum Quaternary sediments 172ndash1725m depth(Rossignol 1964)Etymology Named after Dr Yaakov Ben-Tor (b 1910 ndash d2002) Israeli geologist who was head of the IsraeliGeological Survey at the time of Martine Rossignolrsquos studyand provided her with samplesDistinguishing characters This species typically has a pear-shaped central body with a characteristic apical boss It bearscharacteristic tapering slender gonal and occasionally inter-gonal processes with the two antapical processes being thelongest The process bases may be fenestrate Sutures aremarked by low ridges and sometimes vacuoles are presentin the sutures The wall is smooth to microgranular The cin-gular displacement is relatively large (three times its widthaccording to Reid 1974 one and a half to two times itswidth according to Wall 1965) Tabulation is typical for thegenus according to Wall (1965) and Harland (1968) with fourapical plates although the suture between 1 and 2 is faintand was not observed by Rossignol (1964) or Price ampPospelova (2014) both interpreting this as indicating thepresence of only three apical plates visible on the cyst Thesulcus is often well expressed and straight and widens poster-iorly (Reid 1974) The archeopyle is formed by loss of plate 3rdquoand is reduced with rounded corners (Based on Rossignol1964 p 84ndash85 Harland 1968 p 542ndash543 Reid 1974p 598ndash600 Rochon et al 1999 p 34 Price amp Pospelova2014 p 13 and observations of the participants) MJH notedin draft that specimens from the Last Interglacial of theBaltic have distinctive lateral cingular processes in which thedistal furcations branch abruptly at 90 to the process shafts

and point towards both poles (Head 2007 fig 7jkm)MJH further noted in draft that specimens of S bentorii thathe has observed have archeopyles well defined bythe suturesDimensions Central body length 60ndash73 mm width45ndash63 mm process length 15ndash20 mm (25 mm for the antapi-cal processes) (Rossignol 1964) Central body length58ndash69 mm width 52ndash55 mm length of processes 0ndash20 mm(Reid 1974)Biological affinity Gonyaulax digitale (Pouchet 1883) Kofoid1911 according to Wall amp Dale (1967 p 352) and Dodge(1989 p 283)Intraspecific morphotypes Rossignol (1964) erectedHystrichosphaera bentorii var truncata to encompass speci-mens with shorter processes This variety later was trans-ferred along with the species to Spiniferites (Lentin ampWilliams 1973) Spiniferites bentorii var globus Morzadec-Kerfourn 1979 p 222ndash224 pl 31 fig 10 was introduced toencompass specimens with a spheroidal central body asmaller apical boss and longer more slender processesSeveral subspecies have been described from the Neogenedeposits of the Pannonian Basin in central Europe of whichfour were validly published Spiniferites bentorii subsp buda-jenoensis Seurouto-Szentai 1986 Spiniferites bentorii subsp granu-latus Fuchs amp Seurouto-Szentai 1991 Spiniferites bentorii subspoblongus Seurouto-Szentai 1986 (now Spiniferites oblongusSoliman amp Riding 2017) and Spiniferites bentorii subsp pan-nonicus Seurouto-Szentai 1986 Another four were proposed butnot validly published ldquoSpiniferites bentorii subsp coniunctusrdquoSeurouto-Szentai 1990 ldquoSpiniferites bentorii subsp matraensisrdquo Seurouto-Szentai 1988 ldquoSpiniferites bentorii subsp piriformisrdquo Seurouto-Szentai1988 and ldquoSpiniferites bentorii subsp pseudooblongusrdquo Seurouto-Szentai 1983 None of the eight morphotypes have beenrecorded from PliocenendashQuaternary sedimentsComparison Spiniferites bentorii is similar to Spiniferites lazusin that it can have fenestrate process bases but is distin-guished from Spiniferites lazus by its larger size pear-shape(as opposed to elongate-ovoid) ambitus process form andcingulum displacement (Reid 1974) According to Rochonet al (1999) the central body of Spiniferites bentorii can beovoid but herein we assign such ovoid forms to Spiniferiteslazus Spiniferites bentorii differs from Spiniferites multisphae-rus on the basis of its wall structure which does not containvacuoles unlike Spiniferites multisphaerus (Price amp Pospelova2014) Spiniferites hainanensis has an ovoid central body ForSpiniferites nodosus see Section 226Remarks During the first workshop VP pointed out thatSpiniferites bentorii can have intergonal processes specimensillustrated by Price amp Pospelova (2014) corresponding toSpiniferites bentorii subsp truncatus show such processesRochon et al (1999) also mentioned the presence of occa-sional intergonal processes in this species LL remarked thatthe presence of intergonal processes is not a problem at thespecies level if they are rare or sparse and there is no morethan one between adjacent gonal processes LL furtherremarked that he does not consider the apical boss animportant characteristic but that the pear-shaped centralbody is important Wall amp Dale (1970) while transferring

18 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

processes rough The processes can be perforated due to cor-rosion according to Mao Shaozhi (1989) The processes havewide bases and taper rapidly No sutures are present exceptfor the granules that mark the cingulum dorsally The archeo-pyle is formed by loss of plate 3rdquo (Based on Mao Shaozhi1989 p 139 translated from Chinese by HG and observationsof Achomosphaera sp A of Matsuoka 1983 p 130)Dimensions Central body length (including apical horn)45ndash53 mm width 37ndash45 mm wall thickness 2 mm processlength 10ndash13 mm (Mao Shaozhi 1989)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species can be distinguished from all simi-lar Spiniferites and Achomosphaera species by the uniquedense granules covering the wall and processes its thickouter wall and brown colour (Mao Shaozhi 1989)Remarks Mao Shaozhi (1989) considered Achomosphaeragranulata the same as Achomosphaera sp A of Matsuoka1983 from the Lower to Upper Miocene of the Niigata dis-trict central Japan but we disagree with this assessmentsee also Londeix et al (2018)

The species has been recorded only in Quaternary sedi-ments of the South China Sea by Mao Shaozhi (1989) andMao Shaozhi amp Harland (1993) Londeix et al (2018) consid-ered this species to belong to Hafniasphaera others includ-ing KNM and VP were not convinced because the publishedholotype images are not sharp and the description does notmention the presence of vacuoles

25 Achomosphaera ramosasimilis (Yun Hyesu 1981)Londeix et al 1999

Synonymy Achomosphaera ramulifera subsp ramosasimilisYun Hyesu 1981 p 14ndash15 pl 1 figs 1 8 text-fig 3b

Spiniferites ramuliferus (auct non Deflandre) Reid Reid1974 p 608 pl 4 figs 39ndash40Holotype Yun Hyesu 1981 pl 1 fig 1 text-fig 3b Fensomeet al 1991 figs 1ndash2 p 719 fig 4 p 719 amp 721 [initiallydescribed in German by Yun Hyesu 1981 translated byFensome et al 1991]Type locality Timmermann brickyard new pit near EsbeckWestphalia GermanyType stratum Santonian Upper Cretaceous (Yun Hyesu 1981)Etymology In reference to the close similarity between thisspecies and Spiniferites ramosus (Fensome et al 1991 p 721)Distinguishing characters Thick-walled ovoid to spheroidalcentral body with a granular or smooth pedium and asmooth tegillum bearing hollow distally closed exclusivelygonal processes The processes on the cingulum may be con-nected by crests similar to some processes in the apical andantapical areas There is always a thin apical process occa-sionally with a terminal hooklet Sutures can be partly pre-sent The archeopyle is formed by the loss of plate 3rdquo (Basedon Yun Hyesu 1981 p 14ndash15) and Fensome et al 1991p 719ndash720)Dimensions Central body length 30 (32) 36 mm width 32(39) 42 mm maximum length of processes 16ndash18 mm (YunHyesu 1981)

Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Yun Hyesu (1981) becauseAchomosphaera ramosasimilis possesses a thick wall partialpresence of sutures does not have all processes connectedby crests bears a simple apical process and no antapicalbranched process it differs from Achomosphaera ramuliferaRemarks This species was observed by Londeix et al (1999)from the Lower Pliocene of the Strait of Sicily centralMediterranean Sea LL also considers specimens ofldquoSpiniferites ramuliferusrdquo described by Reid (1974) from recentraised-beach deposits at Woodgrange Northern Ireland UKto be probably Achomosphaera ramosasimilis and notAchomosphaera ramulifera see discussion in Londeix et al(2018) and Gurdebeke et al (2018)

26 Spiniferites alaskensis Marret et al 2001 ex Marretin Fensome amp Williams 2004

Synonymy NoneHolotype Marret et al 2001 pl 1 figs 7ndash9 designated byMarret in Fensome amp Williams (2004)Type locality ODP Site 887 Gulf of Alaska northeasternNorth Pacific Gulf of AlaskaType stratum Marine Isotope Substage 5e Sample 887B-2H-5 65 cm (Marret et al 2001)Etymology Named alaskensis from the type locality (Marretet al 2001)Distinguishing characters Ovoid shape with an apical bossThe cyst wall is thin (lt 1 mm) and has a finely granulate toscabrate surface Processes are gonal and relatively broad(2 mm) each process terminating in a simple trifurcation withpointed ends Low sutural septa are present between proc-esses and define a gonyaulacacean tabulation The archeo-pyle is formed by loss of plate 3rdquo (Based on Marret et al2001 p 384ndash385)Dimensions Central body length 263 (314) 368 mm width236 (293) 315 mm process length 75 (101) 125 mm (Marretet al 2001)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species differs from other Spiniferites spe-cies only by the process shape and termination Spiniferitesalaskensis has broad processes with simple short trifurcatebranches with pointed ends such processes are unusual forlate Quaternary Spiniferites species (Marret et al 2001p 386)Remarks The name was validated by Marret in Fensome ampWilliams (2004) because Marret et al (2001) did not indicatewhich of the illustrations represented the holotype This spe-cies is discussed in Marret amp Mertens (2018)

27 Spiniferites asperulus Matsuoka 1983 (Plate 3figures 1ndash6)

Synonymy NoneHolotype Matsuoka 1983 pl 12 fig 2

PALYNOLOGY 15

Plate 3 1ndash3 Holotype of Spiniferites asperulus Matsuoka 1983 in ventral view at high to low focus 4 Same specimen mid-focus showing wall structure 5 Samespecimen mid-focus showing antapical processes 6ndash8 Holotype of Spiniferites firmus Matsuoka 1983 in ventral view at high to low focus 9 Same specimen show-ing microgranular wall 10 Same specimen showing distal ends of cingular processes Slides provided by KM photographed by KNM All scale bars frac14 10 mmexcept 1ndash3 7ndash9frac14 20 mm

16 K N MERTENS ET AL

Type locality Nishiyama Nishiyama-cho Niigata PrefectureCentral JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin asper (somewhat rough) in ref-erence to the coarsely granular surface of the cyst (Matsuoka1983 p 131)Distinguishing characters The subspheroidal to ovoid cysthas a relatively thick wall The tegillum forms many smallinvaginations in the intratabular area No granules are pre-sent The processes are slender membranous gonal (andoccasionally intergonal) and widen at the base the twoantapical processes are somewhat more robust and con-nected by a slightly elevated crest (Based on Matsuoka1983 p 131ndash132 and observations by KNM)Dimensions Central body length 48ndash69 mm width45ndash64 mm process length up to 16 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison The wall structure is similar to that ofSpiniferites ludhamensis but there are many more invagina-tions in Spiniferites asperulus and Spiniferites ludhamensis issmaller (central body length 38ndash49 mm) and has completelyhollow processes Spiniferites membranaceus has a granu-lar wallRemarks LL expressed the difficulty in differentiating thisspecies from others such as Spiniferites firmus KM stated thatafter having described it back in 1983 he now has no clearrecollection of what this species is LL looked at the holotypebut adds that he did not look at the total population andtherefore finds it hard to say if this is a distinct species or notHe noted that granulation which may occur as a variation inmany species could be a response to specific environmentalconditions The original description mentions the occasionaloccurrence of intergonal processes a feature confirmed byKM This species is discussed in Limoges et al (2018)

28 Spiniferites belerius Reid 1974

Synonymy Non Spiniferites belerius sensu Harland 1977 p98ndash99 pl 1 figs 7ndash10 pl 2 figs 7ndash10 16ndash21 25ndash57 [frac14Spiniferites membranaceus]Holotype Reid 1974 pl 2 figs 12ndash13Type locality Severn Estuary England UKType stratum Modern surface sediments (Reid 1974)Etymology From the Latin belerium which is the Romanname for the promontory on which Landrsquos End CornwallEngland is situated (Reid 1974 p 596)Distinguishing characters The small central body is ovalwith a finely granular wall and an apical boss The processes aregonal and connected by low crests with a typical process shapeThis species was described as having a characteristic largelsquotrumpet-shapedrsquo posterior process at the junction of plates 1and 2 (Reid 1974 p 597 but see Gurdebeke et al 2018)Dimensions Central body length 35ndash42 mm width28ndash37 mm maximum process length 7ndash10 mm posterior pro-cess length 10ndash15 mm (Reid 1974)

Biological affinity Gonyaulax scrippsae Kofoid 1911 asdepicted by Wall amp Dale (1968 p 270 pl 1 fig 14) accord-ing to Reid (1974) LL and KZ noted some similarity to thecyst of Gonyaulax baltica Ellegaard Lewis amp Harding 2002 asproduced in batch-cultures and illustrated by Ellegaard et al(2002) KZ felt she could not understand the cyst-based spe-cies concept here because of the large morphological vari-ation documented in that study VP wondered if anyonecould distinguish cysts of Gonyaulax baltica in the sedimentsME replied that some cysts of Gonyaulax baltica could be thesame as Spiniferites belerius but ideally we should first deriveSpiniferites belerius in culture and compare the phylogeneticsIntraspecific morphotypes NoneComparison LL noted during the first workshop thatSpiniferites bentorii is larger overall and has a hypocyst largerthan the epicyst Spiniferites belerius is similar to Spiniferitesconiconcavus but the latter has wider process basesRemarks During the first and second workshops there wasagreement that this species is problematic LL consideredthis species as having a large morphological variation Healso suggested that this species is defined by central bodysize and shape and the presence of short processes that arenot well developed in the apical region PJM mentioned thatspecimens from the Black Sea show a very wide morpho-logical range and that these could be used as an illustrationof the speciesrsquo various morphologies KNM stated howeverthat potentially different species have been described asSpiniferites belerius given the great variation that he hasobserved globally PJM remarked that this is hard to tellwithout culture studies LL proposed that two categories bedefined for practical purposes typical and atypical morpho-types These categories can then be used in paleoecologicalstudies LL further underlined the wide range in morphologythat appears to characterise many Paratethyan species PJMasked LL whether he considered Spiniferites belerius to beParatethyan and LL answered ldquonordquo He noted that there is aclear regular shape and a wide range of morphological vari-ability and that it is sometimes hard to know if this repre-sents infraspecific variability or more than one species WhenPJM asked what LL considers a Paratethyan species hereplied that it is an ldquoendemicrdquo species with a clearly definedtypical morphology He used Seriliodinium explicatum Eaton1996 Invertocysta sp A of Londeix et al (2009)Invertocysta sp B of Londeix et al (2009) Pterocysta crucifor-mis Rochon et al 2003 Pyxidinopsis psilata (Wall amp Dale inWall et al 1973) Head 1994 Galeacysta etrusca Corradini ampBiffi 1988 and Caspidinium rugosum Marret in Marret et al2004 as good examples of Paratethyan species PJM com-mented in draft that LLrsquos examples include taxa not recordedoutside the Ponto-Caspian basins together with taxa havingwide ranges from the saline western Mediterranean Sea tothe brackish Aral Sea She considers it difficult to justify theword endemic for such a broad grouping of taxa She con-siders that by definition an endemic species means native orrestricted to a certain country or area and should not beused for Paratethyan forms as the Paratethys was a vastEuro-Asian waterway BD suggested that we should decidewhether we can put species into two different groups those

PALYNOLOGY 17

that are stable and well-described and those that originatefrom particular extreme environments that induce increasedecological pressure He said that much of the discussionappears to be focused on forms from extreme environmentsHe noted however that he does not know how to applysuch an approach in practice but proposed that ldquoweirdrdquomorphotypes should be addressed ldquolater onrdquo once clearbasic forms have been established KNM suggested that thetopotype material of Spiniferites belerius should be looked atto evaluate the morphological variability present This hasnow been done by Gurdebeke et al (2018) Limoges et al(2018) have accepted the presence of additional intergonalprocesses in this species

29 Spiniferites bentorii (Rossignol 1964) Wall ampDale 1970

Synonymy Hystrichosphaera bentorii Rossignol 1964 p84ndash85 pl 1 figs 3 3 inset 5ndash8 pl 3 figs 2ndash3 text-figs AndashF

Leptodinium churchillii Harland 1968 p 548 550ndash551 figs12ndash13 22ndash24

non Spiniferites nodosus (Wall 1967) Sarjeant 1970[Synonymy proposed by Reid (1974 p 599) is here rejected]

non Spiniferites bentorii sensu Wall amp Dale 1970 p 52 pl1 figs 26 28Holotype Rossignol 1964 pl 1 figs 3 7ndash8Type locality Ashdod borehole coastal plain IsraelType stratum Quaternary sediments 172ndash1725m depth(Rossignol 1964)Etymology Named after Dr Yaakov Ben-Tor (b 1910 ndash d2002) Israeli geologist who was head of the IsraeliGeological Survey at the time of Martine Rossignolrsquos studyand provided her with samplesDistinguishing characters This species typically has a pear-shaped central body with a characteristic apical boss It bearscharacteristic tapering slender gonal and occasionally inter-gonal processes with the two antapical processes being thelongest The process bases may be fenestrate Sutures aremarked by low ridges and sometimes vacuoles are presentin the sutures The wall is smooth to microgranular The cin-gular displacement is relatively large (three times its widthaccording to Reid 1974 one and a half to two times itswidth according to Wall 1965) Tabulation is typical for thegenus according to Wall (1965) and Harland (1968) with fourapical plates although the suture between 1 and 2 is faintand was not observed by Rossignol (1964) or Price ampPospelova (2014) both interpreting this as indicating thepresence of only three apical plates visible on the cyst Thesulcus is often well expressed and straight and widens poster-iorly (Reid 1974) The archeopyle is formed by loss of plate 3rdquoand is reduced with rounded corners (Based on Rossignol1964 p 84ndash85 Harland 1968 p 542ndash543 Reid 1974p 598ndash600 Rochon et al 1999 p 34 Price amp Pospelova2014 p 13 and observations of the participants) MJH notedin draft that specimens from the Last Interglacial of theBaltic have distinctive lateral cingular processes in which thedistal furcations branch abruptly at 90 to the process shafts

and point towards both poles (Head 2007 fig 7jkm)MJH further noted in draft that specimens of S bentorii thathe has observed have archeopyles well defined bythe suturesDimensions Central body length 60ndash73 mm width45ndash63 mm process length 15ndash20 mm (25 mm for the antapi-cal processes) (Rossignol 1964) Central body length58ndash69 mm width 52ndash55 mm length of processes 0ndash20 mm(Reid 1974)Biological affinity Gonyaulax digitale (Pouchet 1883) Kofoid1911 according to Wall amp Dale (1967 p 352) and Dodge(1989 p 283)Intraspecific morphotypes Rossignol (1964) erectedHystrichosphaera bentorii var truncata to encompass speci-mens with shorter processes This variety later was trans-ferred along with the species to Spiniferites (Lentin ampWilliams 1973) Spiniferites bentorii var globus Morzadec-Kerfourn 1979 p 222ndash224 pl 31 fig 10 was introduced toencompass specimens with a spheroidal central body asmaller apical boss and longer more slender processesSeveral subspecies have been described from the Neogenedeposits of the Pannonian Basin in central Europe of whichfour were validly published Spiniferites bentorii subsp buda-jenoensis Seurouto-Szentai 1986 Spiniferites bentorii subsp granu-latus Fuchs amp Seurouto-Szentai 1991 Spiniferites bentorii subspoblongus Seurouto-Szentai 1986 (now Spiniferites oblongusSoliman amp Riding 2017) and Spiniferites bentorii subsp pan-nonicus Seurouto-Szentai 1986 Another four were proposed butnot validly published ldquoSpiniferites bentorii subsp coniunctusrdquoSeurouto-Szentai 1990 ldquoSpiniferites bentorii subsp matraensisrdquo Seurouto-Szentai 1988 ldquoSpiniferites bentorii subsp piriformisrdquo Seurouto-Szentai1988 and ldquoSpiniferites bentorii subsp pseudooblongusrdquo Seurouto-Szentai 1983 None of the eight morphotypes have beenrecorded from PliocenendashQuaternary sedimentsComparison Spiniferites bentorii is similar to Spiniferites lazusin that it can have fenestrate process bases but is distin-guished from Spiniferites lazus by its larger size pear-shape(as opposed to elongate-ovoid) ambitus process form andcingulum displacement (Reid 1974) According to Rochonet al (1999) the central body of Spiniferites bentorii can beovoid but herein we assign such ovoid forms to Spiniferiteslazus Spiniferites bentorii differs from Spiniferites multisphae-rus on the basis of its wall structure which does not containvacuoles unlike Spiniferites multisphaerus (Price amp Pospelova2014) Spiniferites hainanensis has an ovoid central body ForSpiniferites nodosus see Section 226Remarks During the first workshop VP pointed out thatSpiniferites bentorii can have intergonal processes specimensillustrated by Price amp Pospelova (2014) corresponding toSpiniferites bentorii subsp truncatus show such processesRochon et al (1999) also mentioned the presence of occa-sional intergonal processes in this species LL remarked thatthe presence of intergonal processes is not a problem at thespecies level if they are rare or sparse and there is no morethan one between adjacent gonal processes LL furtherremarked that he does not consider the apical boss animportant characteristic but that the pear-shaped centralbody is important Wall amp Dale (1970) while transferring

18 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Plate 3 1ndash3 Holotype of Spiniferites asperulus Matsuoka 1983 in ventral view at high to low focus 4 Same specimen mid-focus showing wall structure 5 Samespecimen mid-focus showing antapical processes 6ndash8 Holotype of Spiniferites firmus Matsuoka 1983 in ventral view at high to low focus 9 Same specimen show-ing microgranular wall 10 Same specimen showing distal ends of cingular processes Slides provided by KM photographed by KNM All scale bars frac14 10 mmexcept 1ndash3 7ndash9frac14 20 mm

16 K N MERTENS ET AL

Type locality Nishiyama Nishiyama-cho Niigata PrefectureCentral JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin asper (somewhat rough) in ref-erence to the coarsely granular surface of the cyst (Matsuoka1983 p 131)Distinguishing characters The subspheroidal to ovoid cysthas a relatively thick wall The tegillum forms many smallinvaginations in the intratabular area No granules are pre-sent The processes are slender membranous gonal (andoccasionally intergonal) and widen at the base the twoantapical processes are somewhat more robust and con-nected by a slightly elevated crest (Based on Matsuoka1983 p 131ndash132 and observations by KNM)Dimensions Central body length 48ndash69 mm width45ndash64 mm process length up to 16 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison The wall structure is similar to that ofSpiniferites ludhamensis but there are many more invagina-tions in Spiniferites asperulus and Spiniferites ludhamensis issmaller (central body length 38ndash49 mm) and has completelyhollow processes Spiniferites membranaceus has a granu-lar wallRemarks LL expressed the difficulty in differentiating thisspecies from others such as Spiniferites firmus KM stated thatafter having described it back in 1983 he now has no clearrecollection of what this species is LL looked at the holotypebut adds that he did not look at the total population andtherefore finds it hard to say if this is a distinct species or notHe noted that granulation which may occur as a variation inmany species could be a response to specific environmentalconditions The original description mentions the occasionaloccurrence of intergonal processes a feature confirmed byKM This species is discussed in Limoges et al (2018)

28 Spiniferites belerius Reid 1974

Synonymy Non Spiniferites belerius sensu Harland 1977 p98ndash99 pl 1 figs 7ndash10 pl 2 figs 7ndash10 16ndash21 25ndash57 [frac14Spiniferites membranaceus]Holotype Reid 1974 pl 2 figs 12ndash13Type locality Severn Estuary England UKType stratum Modern surface sediments (Reid 1974)Etymology From the Latin belerium which is the Romanname for the promontory on which Landrsquos End CornwallEngland is situated (Reid 1974 p 596)Distinguishing characters The small central body is ovalwith a finely granular wall and an apical boss The processes aregonal and connected by low crests with a typical process shapeThis species was described as having a characteristic largelsquotrumpet-shapedrsquo posterior process at the junction of plates 1and 2 (Reid 1974 p 597 but see Gurdebeke et al 2018)Dimensions Central body length 35ndash42 mm width28ndash37 mm maximum process length 7ndash10 mm posterior pro-cess length 10ndash15 mm (Reid 1974)

Biological affinity Gonyaulax scrippsae Kofoid 1911 asdepicted by Wall amp Dale (1968 p 270 pl 1 fig 14) accord-ing to Reid (1974) LL and KZ noted some similarity to thecyst of Gonyaulax baltica Ellegaard Lewis amp Harding 2002 asproduced in batch-cultures and illustrated by Ellegaard et al(2002) KZ felt she could not understand the cyst-based spe-cies concept here because of the large morphological vari-ation documented in that study VP wondered if anyonecould distinguish cysts of Gonyaulax baltica in the sedimentsME replied that some cysts of Gonyaulax baltica could be thesame as Spiniferites belerius but ideally we should first deriveSpiniferites belerius in culture and compare the phylogeneticsIntraspecific morphotypes NoneComparison LL noted during the first workshop thatSpiniferites bentorii is larger overall and has a hypocyst largerthan the epicyst Spiniferites belerius is similar to Spiniferitesconiconcavus but the latter has wider process basesRemarks During the first and second workshops there wasagreement that this species is problematic LL consideredthis species as having a large morphological variation Healso suggested that this species is defined by central bodysize and shape and the presence of short processes that arenot well developed in the apical region PJM mentioned thatspecimens from the Black Sea show a very wide morpho-logical range and that these could be used as an illustrationof the speciesrsquo various morphologies KNM stated howeverthat potentially different species have been described asSpiniferites belerius given the great variation that he hasobserved globally PJM remarked that this is hard to tellwithout culture studies LL proposed that two categories bedefined for practical purposes typical and atypical morpho-types These categories can then be used in paleoecologicalstudies LL further underlined the wide range in morphologythat appears to characterise many Paratethyan species PJMasked LL whether he considered Spiniferites belerius to beParatethyan and LL answered ldquonordquo He noted that there is aclear regular shape and a wide range of morphological vari-ability and that it is sometimes hard to know if this repre-sents infraspecific variability or more than one species WhenPJM asked what LL considers a Paratethyan species hereplied that it is an ldquoendemicrdquo species with a clearly definedtypical morphology He used Seriliodinium explicatum Eaton1996 Invertocysta sp A of Londeix et al (2009)Invertocysta sp B of Londeix et al (2009) Pterocysta crucifor-mis Rochon et al 2003 Pyxidinopsis psilata (Wall amp Dale inWall et al 1973) Head 1994 Galeacysta etrusca Corradini ampBiffi 1988 and Caspidinium rugosum Marret in Marret et al2004 as good examples of Paratethyan species PJM com-mented in draft that LLrsquos examples include taxa not recordedoutside the Ponto-Caspian basins together with taxa havingwide ranges from the saline western Mediterranean Sea tothe brackish Aral Sea She considers it difficult to justify theword endemic for such a broad grouping of taxa She con-siders that by definition an endemic species means native orrestricted to a certain country or area and should not beused for Paratethyan forms as the Paratethys was a vastEuro-Asian waterway BD suggested that we should decidewhether we can put species into two different groups those

PALYNOLOGY 17

that are stable and well-described and those that originatefrom particular extreme environments that induce increasedecological pressure He said that much of the discussionappears to be focused on forms from extreme environmentsHe noted however that he does not know how to applysuch an approach in practice but proposed that ldquoweirdrdquomorphotypes should be addressed ldquolater onrdquo once clearbasic forms have been established KNM suggested that thetopotype material of Spiniferites belerius should be looked atto evaluate the morphological variability present This hasnow been done by Gurdebeke et al (2018) Limoges et al(2018) have accepted the presence of additional intergonalprocesses in this species

29 Spiniferites bentorii (Rossignol 1964) Wall ampDale 1970

Synonymy Hystrichosphaera bentorii Rossignol 1964 p84ndash85 pl 1 figs 3 3 inset 5ndash8 pl 3 figs 2ndash3 text-figs AndashF

Leptodinium churchillii Harland 1968 p 548 550ndash551 figs12ndash13 22ndash24

non Spiniferites nodosus (Wall 1967) Sarjeant 1970[Synonymy proposed by Reid (1974 p 599) is here rejected]

non Spiniferites bentorii sensu Wall amp Dale 1970 p 52 pl1 figs 26 28Holotype Rossignol 1964 pl 1 figs 3 7ndash8Type locality Ashdod borehole coastal plain IsraelType stratum Quaternary sediments 172ndash1725m depth(Rossignol 1964)Etymology Named after Dr Yaakov Ben-Tor (b 1910 ndash d2002) Israeli geologist who was head of the IsraeliGeological Survey at the time of Martine Rossignolrsquos studyand provided her with samplesDistinguishing characters This species typically has a pear-shaped central body with a characteristic apical boss It bearscharacteristic tapering slender gonal and occasionally inter-gonal processes with the two antapical processes being thelongest The process bases may be fenestrate Sutures aremarked by low ridges and sometimes vacuoles are presentin the sutures The wall is smooth to microgranular The cin-gular displacement is relatively large (three times its widthaccording to Reid 1974 one and a half to two times itswidth according to Wall 1965) Tabulation is typical for thegenus according to Wall (1965) and Harland (1968) with fourapical plates although the suture between 1 and 2 is faintand was not observed by Rossignol (1964) or Price ampPospelova (2014) both interpreting this as indicating thepresence of only three apical plates visible on the cyst Thesulcus is often well expressed and straight and widens poster-iorly (Reid 1974) The archeopyle is formed by loss of plate 3rdquoand is reduced with rounded corners (Based on Rossignol1964 p 84ndash85 Harland 1968 p 542ndash543 Reid 1974p 598ndash600 Rochon et al 1999 p 34 Price amp Pospelova2014 p 13 and observations of the participants) MJH notedin draft that specimens from the Last Interglacial of theBaltic have distinctive lateral cingular processes in which thedistal furcations branch abruptly at 90 to the process shafts

and point towards both poles (Head 2007 fig 7jkm)MJH further noted in draft that specimens of S bentorii thathe has observed have archeopyles well defined bythe suturesDimensions Central body length 60ndash73 mm width45ndash63 mm process length 15ndash20 mm (25 mm for the antapi-cal processes) (Rossignol 1964) Central body length58ndash69 mm width 52ndash55 mm length of processes 0ndash20 mm(Reid 1974)Biological affinity Gonyaulax digitale (Pouchet 1883) Kofoid1911 according to Wall amp Dale (1967 p 352) and Dodge(1989 p 283)Intraspecific morphotypes Rossignol (1964) erectedHystrichosphaera bentorii var truncata to encompass speci-mens with shorter processes This variety later was trans-ferred along with the species to Spiniferites (Lentin ampWilliams 1973) Spiniferites bentorii var globus Morzadec-Kerfourn 1979 p 222ndash224 pl 31 fig 10 was introduced toencompass specimens with a spheroidal central body asmaller apical boss and longer more slender processesSeveral subspecies have been described from the Neogenedeposits of the Pannonian Basin in central Europe of whichfour were validly published Spiniferites bentorii subsp buda-jenoensis Seurouto-Szentai 1986 Spiniferites bentorii subsp granu-latus Fuchs amp Seurouto-Szentai 1991 Spiniferites bentorii subspoblongus Seurouto-Szentai 1986 (now Spiniferites oblongusSoliman amp Riding 2017) and Spiniferites bentorii subsp pan-nonicus Seurouto-Szentai 1986 Another four were proposed butnot validly published ldquoSpiniferites bentorii subsp coniunctusrdquoSeurouto-Szentai 1990 ldquoSpiniferites bentorii subsp matraensisrdquo Seurouto-Szentai 1988 ldquoSpiniferites bentorii subsp piriformisrdquo Seurouto-Szentai1988 and ldquoSpiniferites bentorii subsp pseudooblongusrdquo Seurouto-Szentai 1983 None of the eight morphotypes have beenrecorded from PliocenendashQuaternary sedimentsComparison Spiniferites bentorii is similar to Spiniferites lazusin that it can have fenestrate process bases but is distin-guished from Spiniferites lazus by its larger size pear-shape(as opposed to elongate-ovoid) ambitus process form andcingulum displacement (Reid 1974) According to Rochonet al (1999) the central body of Spiniferites bentorii can beovoid but herein we assign such ovoid forms to Spiniferiteslazus Spiniferites bentorii differs from Spiniferites multisphae-rus on the basis of its wall structure which does not containvacuoles unlike Spiniferites multisphaerus (Price amp Pospelova2014) Spiniferites hainanensis has an ovoid central body ForSpiniferites nodosus see Section 226Remarks During the first workshop VP pointed out thatSpiniferites bentorii can have intergonal processes specimensillustrated by Price amp Pospelova (2014) corresponding toSpiniferites bentorii subsp truncatus show such processesRochon et al (1999) also mentioned the presence of occa-sional intergonal processes in this species LL remarked thatthe presence of intergonal processes is not a problem at thespecies level if they are rare or sparse and there is no morethan one between adjacent gonal processes LL furtherremarked that he does not consider the apical boss animportant characteristic but that the pear-shaped centralbody is important Wall amp Dale (1970) while transferring

18 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Type locality Nishiyama Nishiyama-cho Niigata PrefectureCentral JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin asper (somewhat rough) in ref-erence to the coarsely granular surface of the cyst (Matsuoka1983 p 131)Distinguishing characters The subspheroidal to ovoid cysthas a relatively thick wall The tegillum forms many smallinvaginations in the intratabular area No granules are pre-sent The processes are slender membranous gonal (andoccasionally intergonal) and widen at the base the twoantapical processes are somewhat more robust and con-nected by a slightly elevated crest (Based on Matsuoka1983 p 131ndash132 and observations by KNM)Dimensions Central body length 48ndash69 mm width45ndash64 mm process length up to 16 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison The wall structure is similar to that ofSpiniferites ludhamensis but there are many more invagina-tions in Spiniferites asperulus and Spiniferites ludhamensis issmaller (central body length 38ndash49 mm) and has completelyhollow processes Spiniferites membranaceus has a granu-lar wallRemarks LL expressed the difficulty in differentiating thisspecies from others such as Spiniferites firmus KM stated thatafter having described it back in 1983 he now has no clearrecollection of what this species is LL looked at the holotypebut adds that he did not look at the total population andtherefore finds it hard to say if this is a distinct species or notHe noted that granulation which may occur as a variation inmany species could be a response to specific environmentalconditions The original description mentions the occasionaloccurrence of intergonal processes a feature confirmed byKM This species is discussed in Limoges et al (2018)

28 Spiniferites belerius Reid 1974

Synonymy Non Spiniferites belerius sensu Harland 1977 p98ndash99 pl 1 figs 7ndash10 pl 2 figs 7ndash10 16ndash21 25ndash57 [frac14Spiniferites membranaceus]Holotype Reid 1974 pl 2 figs 12ndash13Type locality Severn Estuary England UKType stratum Modern surface sediments (Reid 1974)Etymology From the Latin belerium which is the Romanname for the promontory on which Landrsquos End CornwallEngland is situated (Reid 1974 p 596)Distinguishing characters The small central body is ovalwith a finely granular wall and an apical boss The processes aregonal and connected by low crests with a typical process shapeThis species was described as having a characteristic largelsquotrumpet-shapedrsquo posterior process at the junction of plates 1and 2 (Reid 1974 p 597 but see Gurdebeke et al 2018)Dimensions Central body length 35ndash42 mm width28ndash37 mm maximum process length 7ndash10 mm posterior pro-cess length 10ndash15 mm (Reid 1974)

Biological affinity Gonyaulax scrippsae Kofoid 1911 asdepicted by Wall amp Dale (1968 p 270 pl 1 fig 14) accord-ing to Reid (1974) LL and KZ noted some similarity to thecyst of Gonyaulax baltica Ellegaard Lewis amp Harding 2002 asproduced in batch-cultures and illustrated by Ellegaard et al(2002) KZ felt she could not understand the cyst-based spe-cies concept here because of the large morphological vari-ation documented in that study VP wondered if anyonecould distinguish cysts of Gonyaulax baltica in the sedimentsME replied that some cysts of Gonyaulax baltica could be thesame as Spiniferites belerius but ideally we should first deriveSpiniferites belerius in culture and compare the phylogeneticsIntraspecific morphotypes NoneComparison LL noted during the first workshop thatSpiniferites bentorii is larger overall and has a hypocyst largerthan the epicyst Spiniferites belerius is similar to Spiniferitesconiconcavus but the latter has wider process basesRemarks During the first and second workshops there wasagreement that this species is problematic LL consideredthis species as having a large morphological variation Healso suggested that this species is defined by central bodysize and shape and the presence of short processes that arenot well developed in the apical region PJM mentioned thatspecimens from the Black Sea show a very wide morpho-logical range and that these could be used as an illustrationof the speciesrsquo various morphologies KNM stated howeverthat potentially different species have been described asSpiniferites belerius given the great variation that he hasobserved globally PJM remarked that this is hard to tellwithout culture studies LL proposed that two categories bedefined for practical purposes typical and atypical morpho-types These categories can then be used in paleoecologicalstudies LL further underlined the wide range in morphologythat appears to characterise many Paratethyan species PJMasked LL whether he considered Spiniferites belerius to beParatethyan and LL answered ldquonordquo He noted that there is aclear regular shape and a wide range of morphological vari-ability and that it is sometimes hard to know if this repre-sents infraspecific variability or more than one species WhenPJM asked what LL considers a Paratethyan species hereplied that it is an ldquoendemicrdquo species with a clearly definedtypical morphology He used Seriliodinium explicatum Eaton1996 Invertocysta sp A of Londeix et al (2009)Invertocysta sp B of Londeix et al (2009) Pterocysta crucifor-mis Rochon et al 2003 Pyxidinopsis psilata (Wall amp Dale inWall et al 1973) Head 1994 Galeacysta etrusca Corradini ampBiffi 1988 and Caspidinium rugosum Marret in Marret et al2004 as good examples of Paratethyan species PJM com-mented in draft that LLrsquos examples include taxa not recordedoutside the Ponto-Caspian basins together with taxa havingwide ranges from the saline western Mediterranean Sea tothe brackish Aral Sea She considers it difficult to justify theword endemic for such a broad grouping of taxa She con-siders that by definition an endemic species means native orrestricted to a certain country or area and should not beused for Paratethyan forms as the Paratethys was a vastEuro-Asian waterway BD suggested that we should decidewhether we can put species into two different groups those

PALYNOLOGY 17

that are stable and well-described and those that originatefrom particular extreme environments that induce increasedecological pressure He said that much of the discussionappears to be focused on forms from extreme environmentsHe noted however that he does not know how to applysuch an approach in practice but proposed that ldquoweirdrdquomorphotypes should be addressed ldquolater onrdquo once clearbasic forms have been established KNM suggested that thetopotype material of Spiniferites belerius should be looked atto evaluate the morphological variability present This hasnow been done by Gurdebeke et al (2018) Limoges et al(2018) have accepted the presence of additional intergonalprocesses in this species

29 Spiniferites bentorii (Rossignol 1964) Wall ampDale 1970

Synonymy Hystrichosphaera bentorii Rossignol 1964 p84ndash85 pl 1 figs 3 3 inset 5ndash8 pl 3 figs 2ndash3 text-figs AndashF

Leptodinium churchillii Harland 1968 p 548 550ndash551 figs12ndash13 22ndash24

non Spiniferites nodosus (Wall 1967) Sarjeant 1970[Synonymy proposed by Reid (1974 p 599) is here rejected]

non Spiniferites bentorii sensu Wall amp Dale 1970 p 52 pl1 figs 26 28Holotype Rossignol 1964 pl 1 figs 3 7ndash8Type locality Ashdod borehole coastal plain IsraelType stratum Quaternary sediments 172ndash1725m depth(Rossignol 1964)Etymology Named after Dr Yaakov Ben-Tor (b 1910 ndash d2002) Israeli geologist who was head of the IsraeliGeological Survey at the time of Martine Rossignolrsquos studyand provided her with samplesDistinguishing characters This species typically has a pear-shaped central body with a characteristic apical boss It bearscharacteristic tapering slender gonal and occasionally inter-gonal processes with the two antapical processes being thelongest The process bases may be fenestrate Sutures aremarked by low ridges and sometimes vacuoles are presentin the sutures The wall is smooth to microgranular The cin-gular displacement is relatively large (three times its widthaccording to Reid 1974 one and a half to two times itswidth according to Wall 1965) Tabulation is typical for thegenus according to Wall (1965) and Harland (1968) with fourapical plates although the suture between 1 and 2 is faintand was not observed by Rossignol (1964) or Price ampPospelova (2014) both interpreting this as indicating thepresence of only three apical plates visible on the cyst Thesulcus is often well expressed and straight and widens poster-iorly (Reid 1974) The archeopyle is formed by loss of plate 3rdquoand is reduced with rounded corners (Based on Rossignol1964 p 84ndash85 Harland 1968 p 542ndash543 Reid 1974p 598ndash600 Rochon et al 1999 p 34 Price amp Pospelova2014 p 13 and observations of the participants) MJH notedin draft that specimens from the Last Interglacial of theBaltic have distinctive lateral cingular processes in which thedistal furcations branch abruptly at 90 to the process shafts

and point towards both poles (Head 2007 fig 7jkm)MJH further noted in draft that specimens of S bentorii thathe has observed have archeopyles well defined bythe suturesDimensions Central body length 60ndash73 mm width45ndash63 mm process length 15ndash20 mm (25 mm for the antapi-cal processes) (Rossignol 1964) Central body length58ndash69 mm width 52ndash55 mm length of processes 0ndash20 mm(Reid 1974)Biological affinity Gonyaulax digitale (Pouchet 1883) Kofoid1911 according to Wall amp Dale (1967 p 352) and Dodge(1989 p 283)Intraspecific morphotypes Rossignol (1964) erectedHystrichosphaera bentorii var truncata to encompass speci-mens with shorter processes This variety later was trans-ferred along with the species to Spiniferites (Lentin ampWilliams 1973) Spiniferites bentorii var globus Morzadec-Kerfourn 1979 p 222ndash224 pl 31 fig 10 was introduced toencompass specimens with a spheroidal central body asmaller apical boss and longer more slender processesSeveral subspecies have been described from the Neogenedeposits of the Pannonian Basin in central Europe of whichfour were validly published Spiniferites bentorii subsp buda-jenoensis Seurouto-Szentai 1986 Spiniferites bentorii subsp granu-latus Fuchs amp Seurouto-Szentai 1991 Spiniferites bentorii subspoblongus Seurouto-Szentai 1986 (now Spiniferites oblongusSoliman amp Riding 2017) and Spiniferites bentorii subsp pan-nonicus Seurouto-Szentai 1986 Another four were proposed butnot validly published ldquoSpiniferites bentorii subsp coniunctusrdquoSeurouto-Szentai 1990 ldquoSpiniferites bentorii subsp matraensisrdquo Seurouto-Szentai 1988 ldquoSpiniferites bentorii subsp piriformisrdquo Seurouto-Szentai1988 and ldquoSpiniferites bentorii subsp pseudooblongusrdquo Seurouto-Szentai 1983 None of the eight morphotypes have beenrecorded from PliocenendashQuaternary sedimentsComparison Spiniferites bentorii is similar to Spiniferites lazusin that it can have fenestrate process bases but is distin-guished from Spiniferites lazus by its larger size pear-shape(as opposed to elongate-ovoid) ambitus process form andcingulum displacement (Reid 1974) According to Rochonet al (1999) the central body of Spiniferites bentorii can beovoid but herein we assign such ovoid forms to Spiniferiteslazus Spiniferites bentorii differs from Spiniferites multisphae-rus on the basis of its wall structure which does not containvacuoles unlike Spiniferites multisphaerus (Price amp Pospelova2014) Spiniferites hainanensis has an ovoid central body ForSpiniferites nodosus see Section 226Remarks During the first workshop VP pointed out thatSpiniferites bentorii can have intergonal processes specimensillustrated by Price amp Pospelova (2014) corresponding toSpiniferites bentorii subsp truncatus show such processesRochon et al (1999) also mentioned the presence of occa-sional intergonal processes in this species LL remarked thatthe presence of intergonal processes is not a problem at thespecies level if they are rare or sparse and there is no morethan one between adjacent gonal processes LL furtherremarked that he does not consider the apical boss animportant characteristic but that the pear-shaped centralbody is important Wall amp Dale (1970) while transferring

18 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

that are stable and well-described and those that originatefrom particular extreme environments that induce increasedecological pressure He said that much of the discussionappears to be focused on forms from extreme environmentsHe noted however that he does not know how to applysuch an approach in practice but proposed that ldquoweirdrdquomorphotypes should be addressed ldquolater onrdquo once clearbasic forms have been established KNM suggested that thetopotype material of Spiniferites belerius should be looked atto evaluate the morphological variability present This hasnow been done by Gurdebeke et al (2018) Limoges et al(2018) have accepted the presence of additional intergonalprocesses in this species

29 Spiniferites bentorii (Rossignol 1964) Wall ampDale 1970

Synonymy Hystrichosphaera bentorii Rossignol 1964 p84ndash85 pl 1 figs 3 3 inset 5ndash8 pl 3 figs 2ndash3 text-figs AndashF

Leptodinium churchillii Harland 1968 p 548 550ndash551 figs12ndash13 22ndash24

non Spiniferites nodosus (Wall 1967) Sarjeant 1970[Synonymy proposed by Reid (1974 p 599) is here rejected]

non Spiniferites bentorii sensu Wall amp Dale 1970 p 52 pl1 figs 26 28Holotype Rossignol 1964 pl 1 figs 3 7ndash8Type locality Ashdod borehole coastal plain IsraelType stratum Quaternary sediments 172ndash1725m depth(Rossignol 1964)Etymology Named after Dr Yaakov Ben-Tor (b 1910 ndash d2002) Israeli geologist who was head of the IsraeliGeological Survey at the time of Martine Rossignolrsquos studyand provided her with samplesDistinguishing characters This species typically has a pear-shaped central body with a characteristic apical boss It bearscharacteristic tapering slender gonal and occasionally inter-gonal processes with the two antapical processes being thelongest The process bases may be fenestrate Sutures aremarked by low ridges and sometimes vacuoles are presentin the sutures The wall is smooth to microgranular The cin-gular displacement is relatively large (three times its widthaccording to Reid 1974 one and a half to two times itswidth according to Wall 1965) Tabulation is typical for thegenus according to Wall (1965) and Harland (1968) with fourapical plates although the suture between 1 and 2 is faintand was not observed by Rossignol (1964) or Price ampPospelova (2014) both interpreting this as indicating thepresence of only three apical plates visible on the cyst Thesulcus is often well expressed and straight and widens poster-iorly (Reid 1974) The archeopyle is formed by loss of plate 3rdquoand is reduced with rounded corners (Based on Rossignol1964 p 84ndash85 Harland 1968 p 542ndash543 Reid 1974p 598ndash600 Rochon et al 1999 p 34 Price amp Pospelova2014 p 13 and observations of the participants) MJH notedin draft that specimens from the Last Interglacial of theBaltic have distinctive lateral cingular processes in which thedistal furcations branch abruptly at 90 to the process shafts

and point towards both poles (Head 2007 fig 7jkm)MJH further noted in draft that specimens of S bentorii thathe has observed have archeopyles well defined bythe suturesDimensions Central body length 60ndash73 mm width45ndash63 mm process length 15ndash20 mm (25 mm for the antapi-cal processes) (Rossignol 1964) Central body length58ndash69 mm width 52ndash55 mm length of processes 0ndash20 mm(Reid 1974)Biological affinity Gonyaulax digitale (Pouchet 1883) Kofoid1911 according to Wall amp Dale (1967 p 352) and Dodge(1989 p 283)Intraspecific morphotypes Rossignol (1964) erectedHystrichosphaera bentorii var truncata to encompass speci-mens with shorter processes This variety later was trans-ferred along with the species to Spiniferites (Lentin ampWilliams 1973) Spiniferites bentorii var globus Morzadec-Kerfourn 1979 p 222ndash224 pl 31 fig 10 was introduced toencompass specimens with a spheroidal central body asmaller apical boss and longer more slender processesSeveral subspecies have been described from the Neogenedeposits of the Pannonian Basin in central Europe of whichfour were validly published Spiniferites bentorii subsp buda-jenoensis Seurouto-Szentai 1986 Spiniferites bentorii subsp granu-latus Fuchs amp Seurouto-Szentai 1991 Spiniferites bentorii subspoblongus Seurouto-Szentai 1986 (now Spiniferites oblongusSoliman amp Riding 2017) and Spiniferites bentorii subsp pan-nonicus Seurouto-Szentai 1986 Another four were proposed butnot validly published ldquoSpiniferites bentorii subsp coniunctusrdquoSeurouto-Szentai 1990 ldquoSpiniferites bentorii subsp matraensisrdquo Seurouto-Szentai 1988 ldquoSpiniferites bentorii subsp piriformisrdquo Seurouto-Szentai1988 and ldquoSpiniferites bentorii subsp pseudooblongusrdquo Seurouto-Szentai 1983 None of the eight morphotypes have beenrecorded from PliocenendashQuaternary sedimentsComparison Spiniferites bentorii is similar to Spiniferites lazusin that it can have fenestrate process bases but is distin-guished from Spiniferites lazus by its larger size pear-shape(as opposed to elongate-ovoid) ambitus process form andcingulum displacement (Reid 1974) According to Rochonet al (1999) the central body of Spiniferites bentorii can beovoid but herein we assign such ovoid forms to Spiniferiteslazus Spiniferites bentorii differs from Spiniferites multisphae-rus on the basis of its wall structure which does not containvacuoles unlike Spiniferites multisphaerus (Price amp Pospelova2014) Spiniferites hainanensis has an ovoid central body ForSpiniferites nodosus see Section 226Remarks During the first workshop VP pointed out thatSpiniferites bentorii can have intergonal processes specimensillustrated by Price amp Pospelova (2014) corresponding toSpiniferites bentorii subsp truncatus show such processesRochon et al (1999) also mentioned the presence of occa-sional intergonal processes in this species LL remarked thatthe presence of intergonal processes is not a problem at thespecies level if they are rare or sparse and there is no morethan one between adjacent gonal processes LL furtherremarked that he does not consider the apical boss animportant characteristic but that the pear-shaped centralbody is important Wall amp Dale (1970) while transferring

18 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Hystrichosphaera bentorii Rossignol to the genus Spiniferitesillustrated a morphotype that does not conform to ourunderstanding of the species This morphotype (Wall amp Dale1970 p 52 pl 1 figs 26 28) bears at least two intergonalprocesses between pairs of gonal processes and despite thepresence of an apical boss we consider it equivalent toSpiniferites hyperacanthus KNM remarked that the cyst-defined plate formula originally provided by Rossignol (1964)is incorrect since all specimens she studied show four apicalplates and not three The species is further discussed byLimoges et al (2018) who report the occasional presence ofintergonal processes

210 Spiniferites bulloideus sensu Wall (1965)

Synonymy Hystrichosphaera bulloidea auct non Deflandre ampCookson 1955 Wall 1965 fig 6 Wall amp Dale 1967 pl 1 figK Wall amp Dale 1968 pl 1 figs 14ndash15 non Spiniferites ramo-sus sensu Wall amp Dale 1970 pl 1 figs 14ndash15 [in contrastwith Harland 1977 p 102] non cyst of Gonyaulax sp affGonyaulax spinifera (frac14Spiniferites ramosum [sic]) in Wall 1971pl 2 fig 4 [in contrast Harland 1977 p 102 considered thisa possible synonym]Holotype Not relevantType locality Not relevant but initially described by Wall(1965) from coastal waters off Woods Hole MA USA (alsofrom the same locality by Wall amp Dale 1967 1968 1970)Type stratum Not relevantEtymology Not relevantDistinguishing characters Ovoid central body without anapical boss and bearing exclusively gonal processes The twoantapical processes are longest membranous and of more orless equal width There may be extensive development ofcrests between the processes The cingulum is displaced Thetabulation is typical for the genus 3ndash4 (suture between 1and 4 faintly visible) 0a 6 6c 5ndash6 1p 1 (Based on Wall1965 p 300ndash302 Wall amp Dale 1968 p 270 and observationsrecorded here)Dimensions Central body length 30ndash40 mm (Wall 1965) and32ndash42 mm (Wall amp Dale 1968) width 28ndash39 mm processlength up to 16 mm (Wall amp Dale 1968)Biological affinity Related to Gonyaulax scrippsae by Wall ampDale (1967 p 352 1968 p 270) Ellegaard et al (2002p 783) recorded cysts of Gonyaulax baltica ldquosimilarto Spiniferites bulloideus sensu Wall amp Dale 1968rdquo in additionto Spiniferites belerius (see remarks therein) The relationshipbetween Gonyaulax baltica and the respective cyst-basedspecies needs further studyIntraspecific morphotypes NoneComparison Spiniferites belerius differs from Spiniferitesbulloideus sensu Wall (1965) in that Spiniferites belerius hasan apical boss Spiniferites bulloideus sensu Wall (1965) dif-fers from Spiniferites falcipedius in being much smallerand from Spiniferites pacificus in not having intergonalprocesses The two antapical tubular processes distinguishthis form from Spiniferites ramosus sensu Rochonet al (1999)

Remarks Specimens Wall (1965) attributed to Spiniferitesbulloideus were recovered from coastal waters off WoodsHole MA USA and later reported by Wall amp Dale (19671968 1970) from the same locality All these studies consid-ered the specimens to belong to Spiniferites bulloideus sensustricto However Harland (1977 p 102) and later Head(1996a p 1205) remarked that ldquoSpiniferites bulloideus sensuWall 1965rdquo (and later referred to by Wall amp Dale 1967 1968)is a distinctive form that should not be attributed to neitherSpiniferites bulloideus sensu stricto nor Spiniferites ramosusSpiniferites bulloideus sensu stricto was first described byDeflandre amp Cookson (1955 as Hystrichosphaera bulloidea pl5 figs 3ndash4 from the Middle Miocene of Balcombe BayVictoria Australia (as mentioned in Deflandre amp Cookson1955 caption to pl 5 figs 3ndash4) It can be described as hav-ing a small spheroidal central body (length 30ndash37 mm) witha thin delicate wall bearing slender processes 10ndash15 mm longthat are probably gonal as well as intergonal (based onDeflandre amp Cookson 1955 p 264 and our observations)The etymology was not specified by the authors but is pre-sumably derived from the Latin bulla (bubble) and the Greekoides (resembling) in reference to the spheroidal centralbody resembling a bubble According to Deflandre ampCookson (1955) Hystrichosphaera bulloidea (now Spiniferitesbulloideus) differs from Hystrichosphaera furcata (now consid-ered a synonym of Spiniferites ramosus) in general outlinedimensions and the nature of the processes and fromHystrichosphaera ramosa (now Spiniferites ramosus) by itsspheroidal central body smaller size and generally moreslender processes During the first round-table discussionparticipants expressed uncertainty as to whether Quaternaryspecimens designated as Spiniferites bulloideus (eg Reid1974 figs 17ndash19 Turon amp Londeix 1988 pl 1 figs 10ndash12McMinn 1991 pl 2 figs 7 12) are conspecific with the holo-type of Deflandre amp Cookson (1955) LL added a note afterthe first workshop that he considered the specimen illus-trated by Turon amp Londeix (1988 pl 1 figs 10ndash12) as a goodexample of Spiniferites bulloideus sensu stricto KNM how-ever remarked that Deflandre amp Cooksonrsquos specimen has adifferent orientation than Turon amp Londeixrsquos (1988) specimenwhich was shown in apical and antapical views so there issome uncertainty as to whether it belongs to Spiniferites bul-loideus since we cannot confirm if the central body is com-pletely spheroidal The same situation is true for imagesshown by McMinn (1991 pl 2 figs 7 12) During the firstround-table discussion there was more or less agreementthat Spiniferites bulloideus sensu stricto could be synonymouswith Spiniferites ramosus possibly as a subspecies The syn-onymy of Spiniferites ramosus with Spiniferites bulloideus waspreviously proposed by Harland (1977) and accepted byMatsuoka (1987a) In either case the holotype should berephotographed but since the type material is in Australiathere was no immediate possibility to do this Either way asBD expressed during his presentation at the second work-shop and followed here Spiniferites bulloideus sensu Wall1965 is different from Spiniferites bulloideus sensu strictobased on the fact that it has two strong antapical processestubular processes and no intergonal processes or

PALYNOLOGY 19

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

membranous developments KNM added that the centralbody of Spiniferites bulloideus sensu Wall 1965 isnot spheroidal

211 Spiniferites coniconcavus De Schepper et al 2004

Synonymy Spiniferites sp 1 of Louwye et al 2004 figs 7rndashtHolotype De Schepper et al 2004 p 628 figs 51ndash520Type locality Verrebroek Dock northern BelgiumType stratum Basal Shelly Unit Lillo Formation upperLower Pliocene (De Schepper et al 2004)Etymology From the Latin conus and concavus in referenceto the principal processes whose bases are cone-shapedwith concave sides (De Schepper et al 2004)Distinguishing characters Broadly ovoid central body bear-ing gonal processes only Process stems are hollow broadand conical with concave sides near the base distallybecoming cylindrical and narrower closed distally with shortand blunt usually trifurcate endings Tabulation expressedby low sutural crests and archeopyle is formed by loss ofplate 3rdquo Operculum is free (Based on De Schepper et al2004 p 628)Dimensions Central body length 38 (399) 40 mm width 34(350) 36 mm process length 7 (98) 125 mm width of processbase 40 (68) 90 mm width of process tip 10 (17) 20 mm(De Schepper et al 2004)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites belerius differs in having an apicalboss Spiniferites bulloideus sensu Wall 1965 has trifurcateprocess terminations that bifurcate distally in contrast to theblunt and reduced process terminations of SpiniferitesconiconcavusRemarks During the first workshop LL wondered how thespecimens illustrated in De Schepper et al (2004 figures517ndash20) differ from Spiniferites belerius SD replied that theprocess bases are wider This species is further discussed byGurdebeke et al (2018)

212 Spiniferites cruciformis Wall and Dale in Wallet al 1973

Synonymy NoneHolotype Wall et al 1973 pl 1 figs 2ndash3Type locality Black Sea Core 1451GType stratum Lower Holocene sediments 665 cm (Wallet al 1973)Etymology From the Latin cruci- (stem of crux cross) and-form in reference to the shape of the central bodyDistinguishing characters Large cysts with characteristiccruciform shape moderately dorsoventrally compressed withsutural septa of variable height that may be perforated (Wallet al 1973 p 21ndash22)Dimensions Central body length 46ndash65 mm width34ndash56 mm depth 28 mm process length up to 28 mm (Wallet al 1973)Biological affinity Unknown

Intraspecific morphotypes Different morphotypes havebeen described by Mudie et al (2001 2018) and Marret et al(2004) In the Caspian Sea morphologies with an elongatedapical boss are included in the species by Marret et al(2004) whereas the holotype and most of the Black Sea mor-phologies have a rounded apexComparison This species is easily distinguishable from allother species of Spiniferites by its flattened cruciform shapewith strongly concave sides above and below the cingulumin equatorial view (Wall et al 1973)Remarks Spiniferites cruciformis was first described by Wall ampDale in Wall et al (1973) from the Lower Holocene of theBlack Sea PJM added a note after the workshop that there isagreement among herself AR LL and Shannon Fergusonthat the species Spiniferites cruciformis Pterocysta cruciformisand Galeacysta etrusca belong to separate genera and donot intergrade AG notes during drafting that he agrees Thisspecies and related taxa are further discussed by Mudie et al(2018) who provide definitions for the terms cruciform gal-eate and pterate as well as a table of characteristics distin-guishing these and other taxa grouped in the so-calledGaleacysta etrusca complex of Popescu et al (2009) MJH indraft provided the following information Although preservedas a single grain mount the holotype is desiccated makingany detailed observations impossible

213 Spiniferites delicatus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 2 figs 20ndash22Type locality Dee Estuary England UKType stratum Modern surface sediment (Reid 1974)Etymology In reference to the delicate sutural membranesjoining the processes (Reid 1974)Distinguishing characters Central body ovoid with micro-granular inner and outer wall layers An apical node or lowboss may be present at the anterior end of 1 and 4Processes are membranous and exclusively gonal with petal-oid process tips High sutural crests connect the processesBoth processes and crests have microgranular surfaces Thecingulum is sinistral and displaced by three times its widthThe tabulation is typical for the genus with an apical seriesof four plates sulcal plates are visible The archeopyle isformed by loss of plate 3rdquo (Based on Reid 1974 p 601ndash602and Rochon et al 1999 p 34)Dimensions Central body length 40ndash60 mm width35ndash54 mm cingulum width 6ndash9 mm maximum process length29 mm (Reid 1974)Biological affinity Unknown but probably Gonyaulax spaccording to Rochon et al (1999)Morphotypes NoneComparison Spiniferites delicatus has processes of similarshape to Spiniferites ristingensis but connected by highsutural crests and a central body wall structure characterisedby a pedium with radial fibers and a thin granular tegillumwhose surface appears microgranular to microreticulateSpiniferites delicatus also differs in having areduced archeopyle

20 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Remarks During the first workshop several participantswondered how to differentiate this species from Spiniferitesristingensis Everybody agreed that the holotype ofSpiniferites delicatus should be reinvestigated LL gave someexamples of specimens he considers typical Spiniferites delica-tus (see Londeix et al 2018) FM stressed the importance ofthe more elaborate development of the crests in Spiniferitesdelicatus There was overall agreement during the secondworkshop that the so-called ldquoskeletal rodsrdquo as first describedby Reid (1974) do not exist and are an optical illusioncreated by the attachment of membranes along the proc-esses This species is further discussed by Gurdebekeet al (2018)

214 Spiniferites elongatus Reid 1974

Synonymy Resting spore of Gonyaulax sp 1 Wall amp Dale1968 pl 1 fig 16 [fide Reid 1974]

cf Hystrichosphaera sp a Harland amp Downie 1969 p 232pl 7 fig 4 [fide Reid 1974]

Spiniferites ellipsoideus Matsuoka 1983 p 132ndash133 pl 13figs 6ndash7

Spiniferites frigidus Harland amp Reid in Harland et al 1980p 213ndash216 fig 2AndashJ

Rottnestia amphicavata Dobell amp Norris in Harland et al1980 p 218ndash220 fig 4AndashN

Rottnestia amphicavata var B Dobell amp Norris in Harlandet al 1980 p 220ndash222 fig 4OndashP fig 8AndashE JndashP

Rottnestia amphicavata var C Dobell amp Norris in Harlandet al 1980 p 222 fig 8FndashI Q R

Spiniferites cf elongatus Harland amp Sharp 1986 pl 1figs 9ndash16Holotype Reid 1974 pl 3 figs 23ndash24Type locality Estuary of the River Ythan Scotland UKType stratum Modern surface sediments (Reid 1974)Etymology From its characteristic elongate shape(Reid 1974)Distinguishing characters Central body elongate andellipsoidal with a smooth to finely microgranulate surfaceand no apical boss Sutural crests are membranous and hol-low with varying height being low around the cingulumand high on the hypocyst and towards the apex Processesare exclusively gonal The cingulum is displaced by lessthan one to two times its width Tabulation is typical forthe genus Sulcal plates are weakly expressed and the sul-cus is aligned parallel to the longitudinal axis increasing tothree times its anterior width posteriorly Plate 600 is triangu-lar long and narrow The archeopyle is formed by loss ofplate 300 and reduced (Based on Reid 1974 p 602ndash603Rochon et al 1999 p 34ndash36 Van Nieuwenhove et al 2018and Gurdebeke et al 2018)Dimensions Central body length 40ndash59 mm width26ndash42 mm wall thickness 08ndash1 mm apical process length6ndash12 mm antapical process length 12ndash16mm lateral processlength 9 mm (Reid 1974)Biological affinity Reid (1974) associated Spiniferites elonga-tus with Gonyaulax scrippsae following observations by Wallamp Dale (1968) Rochon et al (1999) associated it with

Gonyaulax spinifera Ellegaard et al (2003) found the motilestage of Spiniferites elongatus to represent an undescribedspecies of Gonyaulax and using a unified approach to cystand motile stage taxonomynomenclature transferredSpiniferites elongatus to the genus Gonyaulax as G elongata(Reid 1974) Ellegaard et al 2003 The present report followsthe prevailing practice among cyst researchers of using dualtaxonomynomenclature (Head et al 2016 but see Ellegaardet al 2018) and hence the name Spiniferites elongatus ishere retainedIntraspecific morphotypes Spiniferites cf elongatus ofHarland amp Sharp (1986) These cysts were recovered from sur-face sediments of the Norwegian Sea and differ fromSpiniferites elongatus in being ldquosmaller and slightly less mem-branous They are more compact and the processes areshorter and can appear as squat and somewhat lsquodumpyrsquostructures especially on the dorsal surfacerdquo (Harland amp Sharp1986) These forms are now considered part of the morpho-logical spectrum of Spiniferites elongatus and can be infor-mally referred to as Spiniferites elongatus ndash Norwegianmorphotype as noted in Van Nieuwenhove et al (2018)Although the height of sutural crests and their degree ofdevelopment is variable in Spiniferites elongatus they are lesselaborately developed in typical specimens of Spiniferiteselongatus (sensu Reid 1974) than in Rottnestia amphicavataor Spiniferites frigidus and individual processes are moreprominent (Rochon et al 1999) However Van Nieuwenhoveet al (2018) illustrate a morphological continuum and lack ofclear cut-off criteria to distinguish Spiniferites frigidus fromSpiniferites elongatus and that the features of Rottnestiaamphicavata used to place it in this genus can be accommo-dated in Spiniferites Hence Rottnestia amphicavata andSpiniferites frigidus are also considered junior synonyms ofSpiniferites elongatus by Van Nieuwenhove et al (2018) Theyfurther suggest that the ldquoflamboyantrdquo membraneous morph-ology formerly encompassed by Rottnestia amphicavata andSpiniferites frigidus be can informally referred to as Spiniferiteselongatus ndash Beaufort morphotypeComparison Spiniferites elongatus is differentiated from allother species of Spiniferites by its elongate shapeRemarks Spiniferites ellipsoideus was first described byMatsuoka (1983) from Middle to Upper Miocene river cliff sedi-ments of Shinndashshinanogawa Teradomari-cho NiigataPrefecture central Japan During the first workshop partici-pants expressed the opinion that this species is likely a juniorsynonym of Spiniferites elongatus but that the type assemblageshould be checked KM remarked in a personal communicationto KNM that ldquoThis elongate cyst is similar to modern Spiniferiteselongatus but Spiniferites ellipsoideus is smaller than Spiniferiteselongatus and with less development of the antapical mem-brane I think Spiniferites ellipsoideus is an independent speciesfrom Spiniferites elongatusrdquo KNM remarked in draft that meas-urements reported by Matsuoka (1983) (length of 36ndash49mmwidth of cyst 24ndash33mm and length of processes up to 13mm)do not allow an unambiguous differentiation from measure-ments reported for Spiniferites elongatus by Reid (1974) andEllegaard et al (2003) this is confirmed by new measurementsprovided by Van Nieuwenhove et al (2018) who therefore

PALYNOLOGY 21

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

recommend treating Spiniferites ellipsoideus as a junior synonymof Spiniferites elongatus For Spiniferites frigidus and Rottnestiaamphicavata see also Van Nieuwenhove et al (2018)

215 Spiniferites falcipedius Warny amp Wrenn 1997

Synonymy Achomosphaera sp in Head 1997 p 171 figs412ndash416 1510 1511Holotype Warny amp Wrenn 1997 pl 5 figs 1ndash4Type locality Bou Regreg S Core Sale RiffianCorridor MoroccoType stratum Messinian (Warny amp Wrenn 1997)Etymology From the Latin falcipedius meaning bow legs inreference to the wide antapical processes that arise from acommon suturocavate base (Warny amp Wrenn 1997)Distinguishing characters Central body spheroidal toslightly elongate with a microgranular to granular outer wallbearing exclusively gonal large membranous hollow proc-esses The essential criteria for this species are the wideantapical processes connected by a generally high flange(the so-called bow legs) A smaller process may arise fromthe flange connecting the two large antapical processes Thecingulum is offset by a distance equal to the width of thecingulum (Based on Warny amp Wrenn 1997 p 291ndash297)Dimensions Central body length 510ndash748 mm width476ndash646mm process length 102ndash255mm (Warny ampWrenn 1997)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites falcipedius is similar to Spiniferitesmirabilis but differs in having wide exclusively gonal proc-esses (Warny amp Wrenn 1997) During the first workshop LLpointed out the similarity between Spiniferites falcipedius andSpiniferites pacificus although the latter has a much smallercentral body (256ndash268mm) and processes are not as wideand it consistently bears intergonal processes Spiniferitesstrictus is also smaller than Spiniferites falcipedius and hasprocesses that are not as wide Spiniferites membranaceushas a shorter central body a higher flange and a cingulumwhose ends are offset by two cingulum widths MJH addedin draft that Achomosphaera sp of Head (1997) from thePliocene Coralline Crag of eastern England appears similar toSpiniferites falcipedius but is somewhat smaller with a centralbody length of 46ndash55 mm (mean 494mm) (Head 1997) com-pared with 510ndash748 (mean 547 mm) mm for Spiniferites falci-pedius (Warny amp Wrenn 1997) MJH also noted that the distalprocess branches on Spiniferites falcipedius are short wideflat and solid whereas on Spiniferites mirabilis they are longslender and tubular and end in minute bifurcationsRemarks During the first workshop there was a generalcomment that the holotype should be rephotographed butthis has not been possible

216 Spiniferites firmus Matsuoka 1983 (Plate 3figures 7ndash11)

Synonymy None

Holotype Matsuoka 1983 pl 14 figs 5andashcType locality Hachioji Oguni-cho Niigata Prefecture cen-tral JapanType stratum Haizume Formation Lower Pleistocene(Matsuoka 1983)Etymology From the Latin firmus meaning stout in refer-ence to the stout processes (Matsuoka 1983)Distinguishing characters The cyst has a subspheroidal toovoid central body is characterised by a microgranular walland bears exclusively gonal tapering hollow processes withwide bases The processes are connected by low suturesexcept between the two antapical processes where the crestis elevated The archeopyle is formed by loss of plate 3rdquo(Based on Matsuoka 1983 p 134 and observations of theholotype by KM KNM and LL)Dimensions Central body length 40ndash45 mm width38ndash50 mm long processes 16ndash23 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison This species is similar to Spiniferites membra-naceus but differs in having lower crests between theantapical processes and a larger central body It also dif-fers from Spiniferites pachydermus in possessing broadertapering processes with wider bases Spiniferites firmus dif-fers from Spiniferites pseudofurcatus in having a smallercentral body and processes without foliate distal extrem-ities it differs from Spiniferites pseudofurcatus var obliquusin having no processes with a delicate membraneousdistal flare According to Matsuoka (1983) Spiniferitesfirmus is most similar to Spiniferites pacificus andSpiniferites falcipedius but those species have a smallerand a larger central body respectively In additionSpiniferites pacificus has intergonal processes andSpiniferites falcipedius has wider processesRemarks LL indicated in draft that the sutural features ofthis taxon are so faint that it could be considered to belongto Achomosphaera At present it is retained in Spiniferites

217 Spiniferites hainanensis Sun Xuekun amp SongZhichen 1992

Synonymy NoneHolotype Sun Xuekun amp Song Zhichen 1992 pl 1 fig 12Type locality Hainan Island ChinaType stratum Quaternary (Sun Xuekun amp SongZhichen 1992)Etymology Refers to the type locality Hainan IslandDistinguishing characters The central body is slightly ovoidwith a smooth to finely granular wall It bears gonal andintergonal processes connected by low perforated crestsThe archeopyle is formed by the loss of plate 300 (Based onSun Xuekun amp Song Zhichen 1992 p 49)Dimensions Central body length 428ndash490 mm width350ndash420mm process length 105 mm (Sun Xuekun amp SongZhichen 1992)Biological affinity UnknownIntraspecific morphotypes None

22 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Comparison Sun Xuekun amp Song Zhichen (1992) notedthat this species differs from all other known species ofSpiniferites in having uniform perforations at the distalends of the sutural ridges and one or two vacuoles in themiddle part of each process the latter provides the maincriterion for distinguishing this species from SpiniferiteshyperacanthusRemarks This species is discussed further by Limogeset al (2018)

218 Spiniferites hexatypicus Matsuoka 1983 (Plate 4figures 1ndash4)

Synonymy ldquoSpiniferites ovatusrdquo of Bujak 1984 pl 3figs 15ndash18Holotype Matsuoka 1983 pl 13 figs 1andashbType locality Teradomari Teradomarindashcho NiigataPrefecture central JapanType stratum Teradomari Formation Middle to UpperMiocene (Matsuoka 1983)Etymology Derived from the Greek hexathorn typicus (hex-agonal shaped) with reference to the hexagonal cen-tral bodyDistinguishing characters The hexagonal central body lacksan apical boss The wall surface is smooth to finely granularThe processes are exclusively gonal short with simple orsmall bifurcate distal ends (Based on Matsuoka 1983p 133ndash134)Dimensions Central body length 62ndash71 mm width52ndash66 mm process length up to 10 mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Matsuoka (1983) this species issimilar to Spiniferites cingulatus (now Pterodinium cingula-tum) described from the Upper Cretaceous (Senonian) butdiffers in having a hexagonal central body and shorterprocesses The length of the central body distinguishesthis species from Spiniferites ramosus sensu Rochonet al 1999Remarks ldquoSpiniferites ovatusrdquo was invalidly published byBujak (1984) because it is a junior homonym of Spiniferitesovatus Matsuoka 1983) [fide Bujak amp Matsuoka 1986]

Spiniferites hexatypicus was recorded by Matsuoka (1983)from the Middle Miocene to Pliocene of Japan It was alsorecorded by Mao Shaozhi amp Harland (1993) from the UpperPleistocene of the South China Sea All workshop participantsagreed that this is a poorly known species LL considered itto fall within the morphological variability range ofSpiniferites ramosus KNM remarked in draft that the holotypeis very compressed and that the hexagonal shape could bean artefact as a result of compression

219 Spiniferites hyperacanthus (Deflandre amp Cookson1955) Cookson amp Eisenack 1974

Synonymy Hystrichosphaera hyperacantha Deflandre ampCookson 1955 p 264ndash265 pl 6 fig 7 non Spiniferites subspmultiplicatus (Rossignol 1964 p 86 pl 1 fig 14 pl 3 fig 16)

Lentin amp Williams 1973 p 130 [a synonym according toMatsuoka (1985 p 35) but not accepted here]Holotype Deflandre amp Cookson 1955 pl 6 fig 7Type locality Balcombe Bay Victoria AustraliaType stratum Middle Miocene (Deflandre ampCookson 1955)Etymology Not specified by Deflandre amp Cookson (1955)but presumably from the Greek hyper (excess high degree)and akanthos (spine)Distinguishing characters Central body spheroidal with awell-expressed tabulation bearing gonal as well as often twointergonal processes per suture (Based on Deflandre ampCookson 1955 p 264ndash265)Dimensions Central body diameter 54ndash59 mm processlength 13ndash20mm (Deflandre amp Cookson 1955)Biological affinity Motile equivalent Gonyaulax spinifera(Claparede amp Lachmann 1859) Diesing 1866 according toMatsuoka et al (1989 p 94)Intraspecific morphotypes Following Limoges et al (2018)morphotypes with short processes (lt3 mm) should be infor-mally called Spiniferites hyperacanthus ndash short-process mor-photype and specimens with three or more intergonalprocesses between pairs of gonal processes should bereferred to as Spiniferites hyperacanthus ndash multi-intergo-nal morphotypeComparison Spiniferites lenzii Below 1982 was describedfrom the Albian (Lower Cretaceous) of Morocco it has a cen-tral body length of 39ndash44 mm and a width of 38ndash43mm withprocesses of 8ndash15 mm long LL expressed the opinion thatSpiniferites lenzii should be considered a morphotype ofSpiniferites hyperacanthus with higher septa Spiniferites lenziiwas also observed by Matsuoka (1983) from Upper Mioceneto Lower to Middle Pleistocene of the Niigata district (centralJapan) EM noted that Quaternary specimens of Spiniferiteslenzii bear shorter processes than Belowrsquos specimens LL didnot think the specimen depicted in Matsuoka (1983) belongsto this species because Matsuokarsquos specimen shows lowridges rather than elevated septa KNM stated in draft thatMatsuoka (1983) described perforations at the base of theprocesses of his specimens of Spiniferites lenzii which arenot reported by Below (1982) Matsuokarsquos specimensmay correspond to Spiniferites hyperacanthus or anotherspecies For comparison with Spiniferites spinatus seeMatsuoka (1983)Remarks During the first workshop KNM repeated aremark already made by Rossignol (1964) that the antapicalend is not visible on the published micrographs of theholotype of Spiniferites hyperacanthus and therefore wecannot be certain that it has no antapical flange withoutrestudying the holotype Otherwise participants thoughtthis species is well understood KZ considered it as aSpiniferites mirabilis with a reduced antapical flange LLraised the question that if specimens have septa betweenprocesses would they still belong to Spiniferites hyperacan-thus VP stated that the specimens from the continentalslope off Nova Scotia depicted by Rochon et al (1999 pl 7figs 5ndash10) look different from Spiniferites hyperacanthus mdashthese specimens only have one intergonal process per

PALYNOLOGY 23

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Plate 4 1ndash4 Holotype of Spiniferites hexatypicus Matsuoka 1983 in dorsal view at high to low focus Central body length 71 mm Slide provided by KM photo-graphed by KNM 5ndash16 Holotype of Spiniferites ludhamensis Head 1996 in left latero-ventral view at high to low focus Central body length 41 mm Slide providedby MJH photographed by MJH

24 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

suture This species is further discussed by Limoges et al(2018) and Londeix et al (2018)

220 Spiniferites lazus Reid 1974

Synonymy NoneHolotype Reid 1974 pl 3 figs 25ndash27Type locality Pembroke Wales UKType stratum Modern surface sediments (Reid 1974)Etymology From Old French laz (lace) in reference to thefenestrate nature of the process bases (Reid 1974)Distinguishing characters Central body ovoid with a smallapical boss Wall thick with a microgranular to

microreticulate surface Processes are exclusively gonal withwide fenestrate bases and connected by low sutural crestsCingulum displaced by four times its width The archeopyleis formed by loss of plate 3rdquo and is reduced The suturebetween 1 and 4 was not observed by Reid (1974) who indi-cated only three apical plates (Based on Reid 1974 p604ndash605 and Rochon et al 1999 p 36)

Dimensions Central body length 44ndash58 mm width31ndash42 mm thickness 31ndash39 mm wall thickness 1ndash15 mm cin-gulum width 5ndash8 mm process length 12ndash25 mm (Reid 1974)Biological affinity UnknownIntraspecific morphotypes None

Plate 5 1ndash3 Topotype of Spiniferites nodosus (Wall 1967) Sarjeant 1970 in lateral view high to low focus Present width of the central body frac14 46 mm 4ndash7Topotype of Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin amp Williams 1973 in ventral view at high to low focus Central body length 45 mm centralbody width frac14 44 mm Slides provided by David Wall photographed by MJH

PALYNOLOGY 25

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Comparison This species differs from all other Spiniferites onthe basis of its ovoid shape apical boss and fenestrate pro-cess bases Spiniferites bentorii has a pear-shaped centralbody Spiniferites septentrionalis has large fenestrations in the

distal ends of its processes Spiniferites hainanensis has inter-gonal processes (Sun Xuekun amp Song Zhichen 1992)Remarks Reid (1974) mentioned the occurrence of occa-sional intergonal processes in the original description but

Plate 6 1ndash15 Holotype of Spiniferites rhizophorus Head in Head amp Westphal 1999 in ventral view at high to low focus Central body length 50 mm Slide providedby MJH photographed by MJH

26 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Plate 7 1ndash15 Holotype of Spiniferites ristingensis Head 2007 in ventral view at high to low focus Central body length 42 mm Slide provided by MJH photographedby MJH

PALYNOLOGY 27

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Plate 8 1ndash6 Spiniferites scabratus (Wall 1967) Sarjeant 1970 in right lateral view at high to low focus Central body length frac14 56mm central body thickness frac1449mm Slide provided by David Wall photographed by MJH 7ndash9 Holotype of Spiniferites septentrionalis Harland 1979 in ventral view at high to low focus Centralbody length 35mm Photographed by James B Riding 10ndash12 Holotype of Spiniferites strictus Matsuoka 1983 in ventral view at high to low focus Central bodylength 67mm Slide provided by KM photographed by KNM

28 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

this feature has not been observed by anyone attending theworkshops The species is discussed by Gurdebekeet al (2018)

221 Spiniferites ludhamensis head 1996 (Plate 4figures 5ndash16)

Synonymy NoneHolotype Head 1996b fig 12 nos 5ndash9Type locality Royal Society borehole Ludham NorfolkEngland UKType stratum Antian regional pollen zone Gelasian LowerPleistocene (Wall amp Dale 1968 Head 1996b)Etymology Named after the type locality (Head 1996b)Distinguishing characters This species is characterised by acentral body wall with a thin pedium and thicker luxuriaconsisting of a thin tectum supported by funnel-shaped inva-ginations solid at the base where they meet the pediumThe central body is ovoid and has no apical protuberanceThe processes are gonal and occasionally intergonal multi-furcate and bifurcate (possibly with second-order branching)hollow along their length arising from low hollow suturalcrests The sutural crests lack the funnel-shaped invagina-tions The apex is indicated by a tapering spine-like hollowprocess with smaller side branches The archeopyle is formedby loss of plate 3rdquo principal archeopyle suture closely followsplate boundaries and hence has well-defined angles (Basedon Head 1996b p 557)Dimensions Central body length 38 (422) 49mm equatorialdiameter 34 (373) 41 wall thickness 11 (15) 18 mm aver-age process length 10 (129) 15 mm (Head 1996b)Biological affinity UnknownIntraspecific morphotypes None

Comparison This species is distinguished by its unusual wallstructure with numerous invaginations thin-walled hollowprocesses whose branched distal terminations are also hol-low and may form small tubules and sutural folds ratherthan the solid crests which are more typical for the genus(Head 1996b)Remarks During the first workshop EM wondered whetherthere is only one layer with vesicles in Spiniferites ludhamen-sis MJH replied that this is the case EM then stated thatHafniasphaera has several layers and wondered whether thischaracteristic should be used to distinguish Spiniferites andHafniasphaera and if the description of the genus Spiniferitestherefore should be emended KNM noted in draft that theoriginal description of Hafniasphaera describes the wall struc-ture as ldquocomposed of two layers endophragm and peri-phragm One or both of these layers contain numerousevenly distributed vesicles (vacuoles) The vesicles are spher-oidal or if interconnected they may form a fine reticuluminternal in the cyst wallrdquo (Hansen 1977) This species is fur-ther discussed by Limoges et al (2018)

222 Spiniferites membranaceus (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera furcata var membranaceaRossignol 1964 p 86 pl 1 figs 4 9ndash10 pl 3 figs 7 12

Hystrichosphaera ramosa var membranacea (Rossignol1964) Davey amp Williams 1966 p 37

Hystrichosphaera membranacea (Rossignol 1964) Wall1967 p 102Holotype Rossignol 1964 pl 1 figs 4 9ndash10Type locality Ashkelon borehole St 39D coastalplain Israel

Plate 9 1ndash6 Impagidinium inaequalis (Wall and Dale in Wall et al 1973) Londeix et al 2009 in left lateral view at high to low focus central body length frac14 54 mmSlide provided by David Wall photographed by MJH

PALYNOLOGY 29

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Type stratum Pleistocene or Holocene sediments(Rossignol 1964)Etymology Not specified by Rossignol (1964) but presum-ably in reference to the distinct membranous flange in theantapical region that characterises this speciesDistinguishing characters Central body ovoid with noapical boss Surface of the outer wall is microgranulate tomicrorugulate Processes are exclusively gonal and are dis-tally furcate with recurved bifurcate tips Sutural crests aremostly low but are high at the antapex where they form aconspicuous membrane between antapical processes Thecingulum is inclined and displaced by twice its width andthe sulcus is slightly sigmoid and moderately wideTabulation is typical for the genus and sulcal plates are vis-ible The archeopyle is formed by loss of plate 3rdquo (Based onRossignol 1964 p 86 Reid 1974 p 605ndash606 Lewis et al1999 p 115ndash117 Rochon et al 1999 p 36ndash38 Ellegaardet al 2003 p 154ndash156)Dimensions Central body length 57mm width 50mm processlength 20ndash25mm (Rossignol 1964) Central body length34ndash44mm width 34ndash43mm cingulum width 5ndash8mm length ofantapical flange 2ndash21mm process length 12ndash17mm (Reid 1974)Biological affinity Previously Gonyaulax spinifera (Claparedeamp Lachmann 1859) Diesing 1866 according to Dale (1976table 2 p 45) and Dodge (1989 p 289) Ellegaard et al(2003) found the motile stage of Spiniferites membranaceusto represent an undescribed species of Gonyaulax and usinga unified approach to cyst and motile stage taxonomynomenclature transferred Spiniferites membranaceus to thegenus Gonyaulax as G membranacea (Rossignol 1964)Ellegaard et al 2003 However we follow the prevailingpractice among cyst researchers of using dual taxonomynomenclature (Head et al 2016 but see Ellegaard et al2018) and hence the name Spiniferites membranaceus ishere retainedIntraspecific morphotypes LL remarked during the work-shop that specimens with an antapical flange such as thosedepicted by Wall (1967 pl 14 figs 14ndash15) from theCaribbean should be referred to as Spiniferites cf membrana-ceus because overall the processes are relatively short andthe antapical flange is supported by distinctly stout and rod-like processes unlike those of the holotype and specimensmentioned above as typicalComparison Spiniferites membranaceus is distinguished fromSpiniferites mirabilis by its absence of intergonal processesSee Londeix et al (2018) for an extended comparison withother similar speciesRemarks Wall et al (1977) (repeated by Harland 1983)stated that Spiniferites membranaceus sensu Reid (1974) isprobably not conspecific with that of Rossignol (1964) Toverify this KNM proposed that Rossignolrsquos holotype shouldbe restudied but EM replied that it is probably lost MJHsuggested that the topotype material be restudied Rochonet al (1999) noted the presence of occasional intergonalprocesses but this could not be confirmed by workshop par-ticipants This species is further discussed by Gurdebekeet al (2018)

223 Spiniferites mirabilis (Rossignol 1964)Sarjeant 1970

Synonymy Hystrichosphaera mirabilis Rossignol 1964 p86ndash87 pl 2 figs 1ndash3 pl 3 figs 4ndash5 [The name was invalidin Rossignol (1962 p 132) because no illustration was pro-vided and no holotype designated]

Spiniferites splendidus Harland 1979 p 537 pl 3 figs 1ndash2Holotype Rossignol 1964 pl 2 figs 1ndash2Type locality Ashdod Yam borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology From the Latin mirabilis meaning admirable(Rossignol 1964)Distinguishing characters Central body ovoid with asmooth to granular surface The processes are gonal as wellas intergonal slender and connected by generally low tobarely discernible crests However a prominent antapicalflange connects the two antapical processes The sulcus isslightly oblique and the cingulum is displaced by two timesits width The archeopyle is formed by the loss of plate 3rdquoand is reduced (Based on Rossignol 1964 p 86ndash87 and newobservations by AL LL and KNM)Dimensions Central body length 40ndash70 mm width35ndash60 mm process length 15ndash22 mm (Rossignol 1964)Biological affinity Related to Gonyaulax spinifera accordingto cyst-theca experiments by Wall amp Dale (1967 p 352) andWall amp Dale (1968 p 270) reproduced by Dale (1983) andDodge (1985 p 215 1989 p 289) Sonneman amp Hill (1997)and Morquecho et al (2009) also conducted cyst-thecaexperiments on Spiniferites mirabilis and also identified themotile stage as Gonyaulax spiniferaIntraspecific morphotypes Spiniferites mirabilis subsp ser-ratus Limoges et al (2018) originally described byMatsuoka (1983) from the Pliocene (or younger) NishiyamaFormation Ishiji Nishiyama-cho Niigata prefecture (centralJapan) The central body length is 45ndash54 mm width47ndash50 mm with process length 7ndash9 mm It differs fromSpiniferites mirabilis subsp mirabilis (autonym) in havingldquomany short conical to subconical intergonal processesrdquo(Matsuoka 1983) Limoges et al (2018) consider specimenswith three or more intergonals as belonging to Spiniferitesmirabilis subsp serratusComparison The main distinction between this species andSpiniferites hyperacanthus is the presence of an antapicalflange The main distinction between Spiniferites mirabilisand Spiniferites membranaceus is the presence of intergonalprocesses in Spiniferites mirabilisRemarks Spiniferites splendidus was described from theUpper MiocenendashLower Pliocene of DSDP Hole 400 A (Bay ofBiscay) by Harland (1979) According to Harland (1979)Spiniferites splendidus is larger than Spiniferites mirabilis andhas many membranous processes However during theworkshop MJH remarked that Harland (1979) did not providemeasurements of Spiniferites mirabilis but that his illustratedspecimens are not markedly different in size from the singlespecimen of Spiniferites splendidus illustrated by Harland(1979) During the round table discussion LL stated thatthere are fewer processes on Spiniferites splendidus than onSpiniferites mirabilis MJH replied that Harland (1979) did not

30 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

describe the number of processes as diagnostic so maybehe just illustrated an aberrant specimen SD remarked thatthere seem to be membraneous processes on the sulcussomething not seen in Spiniferites mirabilis MJH expressedsome concern about the fact that only two specimens wereillustrated by Harland (1979) and noted a further need tostudy topotype material The holotype was rephotographedby James B Riding and the synonymy is further discussed byLimoges et al and Londeix et al (2018)

LL remarked during the first workshop that it is necessaryto add to the original description that processes are hollowand can be distally open as is the antapical flange Headded that he has however only seen this in Miocene speci-mens AR stated during draft that he has made similar obser-vations in specimens from Argentina VP added that it isremarkable that Spiniferites mirabilis and Spiniferites membra-naceus were described from the same sample LL added afterthe first workshop that several modern Mediterranean speci-mens have hollow and open processes KNM also added thatit is remarkable that this species can have very faint to nosutures which could place it in Achomosphaera The speciesis further discussed by Limoges et al (2018) and Londeixet al (2018)

224 Spiniferites multisphaerus Price ampPospelova 2014

Synonymy NoneHolotype Price amp Pospelova 2014 pl 1 figs 1ndash13Type locality Core MD02ndash2515 Guaymas Basin Gulf ofCalifornia MexicoType stratum Upper Quaternary sediments (Price ampPospelova 2014)Etymology From the Latin multi (many) and sphaerae(spheres) in reference to the characteristic central body wallstructure that features multiple vacuoles (Price ampPospelova 2014)Distinguishing characters Ovoid to pear-shaped centralbody with a pronounced apical protuberance Wall is rela-tively thick and contains a single layer of vacuoles Processesare stubby furcate and also contain vacuoles Tabulation isclearly expressed and is typical of the genus The archeopyleis formed by loss of plate 3rdquo (Based on Price amp Pospelova2014 p 7ndash13)Dimensions Central body length (including apical protuber-ance) 426 (524) 665mm wall thickness 10 (15) 21mmmean length of processes 15 (44) 80mm (Price ampPospelova 2014)Biological affinity UnknownIntraspecific morphotypes NoneComparison Spiniferites multisphaerus is distinguished bythe presence of vacuoles in the cyst wall processes andsutural septa this feature is not known in other QuaternarySpiniferites species In terms of extant taxa Spiniferites multi-sphaerus most closely resembles Spiniferites bentorii as bothhave a pronounced apical boss and lsquoshouldersrsquo on the epi-cyst It is distinguished from Spiniferites bentorii by the pres-ence of the vacuoles In addition the surface of Spiniferites

multisphaerus appears reticulate whereas Spiniferites bentoriiappears smooth to microgranular (Price amp Pospelova 2014)Spiniferites multisphaerus differs from Spiniferites ludhamensisin that the latter has a wall-structure comparable to bub-ble-wrapRemarks During the first round table discussion there wasthe general question put forward by EM as to whether thisspecies should be transferred to Hafniasphaera a genusdescribed by Hansen (1977) that is distinguished fromSpiniferites by the presence of one or two layers containingspheroidal vacuoles or vacuoles forming a fine reticuluminside the cyst wall Since Spiniferites multisphaerus containsa single layer of vacuoles (as pointed out by AP) it could beassigned to Hafniasphaera VP expressed concern that it isdifficult to draw the line with other increasingly more granu-lated walls EM pointed out that if we accept a vacuolatedwall for Spiniferites the genus description of Sarjeant (1970)has to be emended This species is further discussed byLimoges et al (2018) Londeix et al (2018) transferred thisspecies to Hafniasphaera

225 Spiniferites nanus Matsuoka 1976

Synonymy non Spiniferites bulloideus (Deflandre amp Cookson1955) Sarjeant 1970Holotype Matsuoka 1976 pl 28 figs 1ndash3Type locality Nara City JapanType stratum Utahime Member Saho Formation LowerPleistocene (Matsuoka 1976)Etymology Not specified by Matsuoka (1976) but is fromthe Latin nanus for dwarf in reference to the short processlength (confirmed by KM) The epithet is a nounin appositionDistinguishing characters The central body has a roundedconical epicyst and a rounded hypocyst The outer wall issmooth or very finely granular The sulcal region is almoststraight and widens posteriorly The relatively broad cingu-lum is displaced by one to two times its own width Theshort processes with bifurcate trifurcate or acuminate tipsare mostly gonal with intergonal processes occurring occa-sionally The short acuminate processes with wide bases areformed from membraneous sutures The tabulation is typicalfor the genus and the small triangular plate 600 is extendedlongitudinally The archeopyle is formed by loss of plate 300

and is reduced (Based on Matsuoka 1976 p 111)Dimensions Central body length 41ndash55 mm width35ndash54 mm process length 5ndash11 mm (Matsuoka 1976 p 111)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1976) remarked that Spiniferites bul-loideus differs from Spiniferites nanus in having a relativelylarger central body (length 54ndash64 mm width 30ndash37mm) andlonger and exclusively gonal processes (10ndash15mm) measure-ments taken from Deflandre amp Cookson (1955)Remarks KM stated in a communication to KNM in draftthat ldquoI now believe this species is a junior synonym of therelatively broad species concept of Spiniferites bulloideusSpiniferites nanus is a short-process form of Spiniferites

PALYNOLOGY 31

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

bulloideus which presumably exhibits similar morphologicalvariability in processes as documented for cysts ofProtoceratium reticulatum and cysts of Lingulodinium poly-edrardquo However according to the original descriptionSpiniferites nanus can sometimes have intergonal processesThus Spiniferites nanus may not be synonymous withSpiniferites bulloideus but we recommended that the formerspecies be restricted to the holotype Matsuoka (1987b) alsoobserved specimens from the Holocene close to Kawasakicity (Japan) See also Limoges et al (2018)

226 Spiniferites nodosus (Wall 1967) Sarjeant 1970(Plate 5 figures 1ndash3)

Synonymy Hystrichosphaera nodosa Wall 1967 p 101 pl14 figs 7ndash9 text-fig 2 non Spiniferites [as Hystrichosphaera]bentorii according to Reid (1974 p 598) we follow Lentin ampWilliams (1981 p 264) in retaining Spiniferites nodosusHolotype Wall 1967 pl 14 figs 7ndash9Type locality Core A24018 Cariaco Trench off VenezuelaType stratum Quaternary (Wall 1967)Etymology Not specified by Wall (1967) but probably fromthe Latin nodosus meaning knotty in reference to the shapeof the processesDistinguishing characters Ovoid central body with weaklytruncated apices The plate areas are defined by distinctbut low (1 mm) sutural septa and are typical in numberand arrangement for the genus The characteristic proc-esses are exclusively gonal small either bifurcate or trifur-cate and recurve strongly towards their own bases or liealong the cyst surface so that there appears to be a smallpad of tissue at each junction Only rarely do the proc-esses project more than a few microns above the test wallThe archeopyle is formed by loss of plate 3rdquo and has aweakly inclined cingulum (Based on Wall 1967p 101ndash102)Dimensions Central body length 31ndash62 mm width 28ndash52 mm(Wall 1967)Biological affinity Related to Gonyaulax digitale accordingto Wall amp Dale (1967 p 352) repeated in Wall amp Dale (1968p 269)Intraspecific morphotypes NoneComparison PJM remarked that this species is difficult todistinguish from Spiniferites bentorii indeed Reid (1974) hadproposed that Spiniferites nodosus is a junior synonym ofSpiniferites bentorii However LL is not convinced that theholotype of Spiniferites nodosus falls within the range ofSpiniferites bentorii because its central body lacks the typicalpear-shape and is smaller (31ndash62 mm in length 28ndash52mm inwidth) Therefore we recommend restricting this name tothe holotypeRemarks MJH noted during the first workshop that it wasunclear whether one or two specimens were assigned asthe holotype and thus questioned whether the speciesname is valid However in draft a personal communica-tion of KNM with David Wall confirmed that it is a singlespecimen that was rotated into different views Wall (1967p 102) remarked that ldquoThis species gives the impression

that as a cyst it was closely pressed against its parentalthecal covering and that the spines were unable todevelop fully but it is not necessarily an immature formrdquoHowever participants agreed that the ldquostrongly recurvedprocessesrdquo of Spiniferites nodosus are probably due to pres-ervation MJH in draft provided the following informationldquoThe holotype is preserved on a single grain mount and isbroken and somewhat compressed The central body hasan apical protrusion of about 10 mm and a wall thicknessof about 03 mm The surface is very faintly and finelygranulate (scabrate) Sutural septa are about 15 mm highThere is a slight separation of wall layers at the base ofsepta Processes are mostly gonal although one intergonalprocess was clearly identified Gonal process are stronglyshortened by crumpling making it impossible to deter-mine their exact morphology I agree that the name isbest restricted to the holotyperdquo

227 Spiniferites pachydermus (Rossignol 1964)Reid 1974

Synonymy Hystrichosphaera furcata var pachydermaRossignol 1964 p 86 pl 1 figs 1ndash2 pl 3 fig 6

ldquoHystrichosphaera ramosa var pachydermardquo Harland ampDownie 1969 p 232

Spiniferites ramosus subsp pachydermus (Rossignol 1964)Lentin amp Williams 1973 p 130Holotype Rossignol 1964 pl 1 figs 1ndash2Type locality Ashdod borehole 150 coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Derived from the Greek pachy thick and dermaskin (Head 1996a p 1232) in reference to the thickcyst wallDistinguishing characters Central body ovoid with a lowapical boss The wall is thick (gt1 mm) with a microreticulateperforate surface The processes are stout and exclusivelygonal and joined by mostly low sutural septae that expressthe tabulation Notable exceptions to these low crest heightscan be found between the two processes between the anter-ior sulcal plate and the apical plates 1 and 4 which arealways connected by a crest that reaches the distal ends ofthe processes and the elevated crest heights for closelyspaced processes along the cingulum particularly for theprocesses contacting on the left of 1c and 200 0 The processesare variable in length becoming progressively shorter fromthe antapex to the apex The processes are formed at theconvergence of three sutures Distally the processes arepetaloid and trifurcate with slender recurved bifid tips Oftenclaustra are present at the base of the processes or on theconnecting crests Specimens with processes that appearbroken off were also encountered Tabulation is typical forthe genus The antapical plate is asymmetrical The cingulumis sinistral and displaced by a distance 35 times its widthThe sulcus is moderately wide and almost straight and allsulcal plates are visible The archeopyle is formed by loss ofplate 3rdquo is reduced and has a serrated margin The opercu-lum is free (Based on Mertens et al 2015 p 563ndash566)

32 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Dimensions Central body length 61mm width 52 mm wallthickness 3 mm (Rossignol 1964) Central body length33ndash54 mm width 37ndash44 mm cingulum width 5ndash9 mm max-imum process length 19 mm (Reid 1974) Central body length(including low apical boss) 366 (426) 481mm width (meas-ured along the cingulum) 306 (357) 410mm average lengthof six processes per cyst 53 (102) 177 mm (Mertenset al 2015)Biological affinity Gonyaulax ellegaardiae Mertens AydinTakano Yamaguchi amp Matsuoka 2015 according to Mertenset al (2015)Intraspecific morphotypes NoneComparison The microreticulateperforate wall distinguishesSpiniferites pachydermus from all other described species ofSpiniferites (Mertens et al 2015)Remarks The consensus during the first workshop was thata restudy of the holotype was needed but subsequentefforts by EM to locate the holotype at the Museum NationaldrsquoHistoire Naturelle in Paris have not been successful LLidentifies Spiniferites pachydermus through its thick wall andsomewhat larger size than other species However KZ repliedthat Mediterranean species are always larger than elsewhereand that this might complicate identification FS agreed LLremarked that in the specimen illustrated by Rossignol (1964fig 6) the wall is very thick and thus she compared it withAchomosphaera crassipellis (wall thickness 15ndash2 mmDeflandre amp Cookson 1955 p 265) but generally the wall ofSpiniferites pachydermus is not that thick

ldquoHystrichosphaera ramosa var pachydermardquo as proposedby Harland amp Downie (1969) is an invalid combination sincethe basionym was not fully referenced

228 Spiniferites pacificus Zhao Yunyun amp Morzadec-Kerfourn 1994

Synonymy non Spiniferites cf delicatus of Matsuoka 1992pl 1 fig 4

Holotype Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1figs 1andashc

Type locality ODP Leg 126 Site 791 IzundashBonin regionwestern North Pacific

Type stratum Lower or Upper Pleistocene (Zhao Yunyun ampMorzadec-Kerfourn 1994)Etymology Derived from its presence in the Pacific Ocean(Zhao Yunyun amp Morzadec-Kerfourn 1994)Distinguishing characters Central body spheroidal to ovoidwith a microgranular surface and bearing both gonal andintergonal processes (one between each of a pair of gonalprocesses) The two large hollow antapical processes are lon-ger and membranous and open distally The tabulation istypical for the genus 3ndash4 0a 6 6c 5ndash6 1p 1 The archeo-pyle is formed by loss of plate 3rdquo (Based on Zhao Yunyun ampMorzadec-Kerfourn 1994 p 268ndash269)Dimensions Central body length 256ndash353 mm width288ndash299mm average length of gonal processes 108mmaverage length of antapical processes 132 mm (Zhao Yunyunamp Morzadec-Kerfourn 1994)Biological affinity Unknown

Intraspecific morphotypes NoneComparison This species differs from other species thathave two pronounced antapical processes (Spiniferites falcipe-dius Spiniferites firmus and Spiniferites bulloideus sensu Wall1965) by the presence of intergonal processesRemarks Spiniferites cf delicatus of Matsuoka (1992) doesnot have intergonal processes as here confirmed by KM andthus is not conspecific with Spiniferites pacificus

KNM remarked during the first workshop that it was notclear whether the specimens illustrated through light micro-graphs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 1 figs1ndash3) show intergonal processes although specimens illus-trated through SEM micrographs (Zhao Yunyun amp Morzadec-Kerfourn 1994 pl 2 figs 1ndash3) clearly bear intergonal proc-esses LL mentioned in draft that the sutural features of thisspecies are so faint that it could be considered as belongingto Achomosphaera

229 Spiniferites pseudofurcatus subsp obliquus (Wall1967) Lentin amp Williams 1973 (Plate 5figures 4ndash7)

Synonymy Hystrichosphaera tertiaria var obliqua Wall 1967p 103 pl 14 fig 16 text-fig 2Holotype Wall 1967 pl 14 fig 16Type locality Core A254327 Yucatan Basin Caribbean SeaType stratum Upper PleistocenendashHolocene 400 cm depth(Wall 1967)Etymology Not specified by Wall (1967) but the subspecificepithet presumably derives from the oblique nature of thelongitudinal sulcusDistinguishing characters The central body is ovoid some-times with a low apical protuberance and has a smooth toweakly granular surface The processes are distinctive gonaltrifurcate with secondary branches that tend to be paralleland often are connected by delicate membranes occasionalintergonal processes are present Two dorsal antapical proc-esses are prominent The tabulation is typical for the genusThe sulcus is oblique narrows anteriorly and widens poster-iorly the cingulum is strongly displaced at its ends (Basedon Wall 1967 p 103)Dimensions Central body length 40ndash50 mm process length10ndash12 mm (Wall 1967)Biological affinity UnknownIntraspecific morphotypes NoneComparison According to Wall (1967) Spiniferites pseudo-furcatus subsp obliquus (as Hystrichosphaera tertiaria varobliqua) is a small morphotype of Hystrichosphaera tertiariaEisenack amp Gocht 1960 The latter was first described byEisenack (1954 as ldquoHystrichosphaera cf furcatardquo) from theOligocene with a central body length of 50ndash70 mm andtotal diameter of 80ndash133 mm Hystrichosphaera tertiaria isnow considered a synonym of Spiniferites pseudofurcatus(Klumpp 1953) Sarjeant 1970 (Gocht 1969 Sarjeant 1970)the type of which is from the Upper Eocene of GermanySpiniferites bentorii has a more prominent apical protuber-ance (Wall 1967)

PALYNOLOGY 33

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Remarks Wrenn amp Kokinos (1986 pl 7 fig 1ndash2) alsodepicted this subspecies from the Lower to Upper Plioceneof the Gulf of Mexico (image reproduced in de Vernal ampMudie 1992 pl 4 fig 3) During the first workshop MJHnoted that he has seen the holotype and that it does notlook particularly distinctive - he would just call it Spiniferitessp KZ suggested that the subspecies should be restricted tothe holotype LL stated that the assemblages studied byWrenn amp Kokinos (1986) included many reworked specimensso he questions the status of the type material of this taxon

230 Spiniferites ramosus sensu Rochon et al (1999)

Synonymy Spiniferites bulloideus (auct non Deflandre ampCookson) Sarjeant Reid 1974 pl 2 figs 17ndash19

Spiniferites ramosus (auct non Ehrenberg) Mantell Harland1977 pl 1 figs 5ndash6

Spiniferites ramosus (auct non Ehrenberg) Mantell sensulato Rochon et al 1999 pl 9 figs 4ndash6

non Spiniferites ramosus var ramosus (Ehrenberg) MantellDavey amp Rogers 1975 pl 1 fig 5Holotype Not relevantType locality Not relevantType stratum Not relevantEtymology Not relevantDistinguishing characters Central body ovoid to spheroidalwith or without apical boss and a smooth pedium and tegil-lum Processes are exclusively gonal long hollow and havelong distal furcations with bifurcate tips Sutural crests arelow but rise towards the gonal processes to which they con-nect Narrow thread-like pairs of trabeculae between adja-cent process tips are sometimes developed Tabulation istypical for the genus with plates 1 and 4 appearing to befused on the cysts The cingulum is displaced by its ownwidth and the archeopyle is formed by loss of plate 3rdquo(Based on Reid 1974 p 600ndash601 as Spiniferites ramosus inHarland 1977 p 102ndash103 and as Spiniferites ramosus sensulato in Rochon et al 1999 p 38)Dimensions Central body length 30ndash46mm (Rochon et al1999) and 41ndash46mm (Reid 1974) width 19ndash27mm cingulumwidth 6ndash8mm maximum process length 8ndash16mm (Reid 1974)Biological affinity Through incubation experiments onmaterial from Harrington Sound (Bermuda) Wall amp Dale(1970) related Spiniferites ramosus to Gonyaulax spiniferahowever their specimens bear both gonal and occasionalintergonal processes They later called it the ldquoBermuda typerdquoor ldquoSpiniferites ramosus sensu Wall amp Dale 1970rdquo (Wall et al1977) The lack of detail in the description particularly of thecyst wall does not exclude the possibility that this is a differ-ent taxon from Spiniferites ramosus sensu lato of Rochonet al (1999) Much later Lewis et al (1999) relatedSpiniferites ramosus to Gonyaulax spinifera with the morph-ology of the cyst similar to the morphological conceptdescribed above Cysts attributed to Spiniferites ramosusformed in culture experiments and related to Gonyaulax spi-nifera by Rochon et al (2009) correspond relatively well tothe description of Rochon et al (1999) but showed a widerrange of surface ornamentation of the tegillum varying from

shagreenate to granulate and exhibiting occasional develop-ment of trabeculae joining the process tipsIntraspecific morphotypes NoneComparison Spiniferites ramosus sensu lato of Rochon et al(1999) differs from Spiniferites bulloideus sensu Wall (1965)primarily in its uniform slender processes the latter havingtwo membranous stout antapical processesRemarks Spiniferites ramosus the type of Spiniferites wasfirst described by Ehrenberg (1837 as Xanthidium ramosum)from the Upper Cretaceous (Senonian) of GermanyEhrenberg did not designate a holotype and provided draw-ings without a description Davey amp Williams (1966 p 33ndash34)emended the description of Spiniferites and designatedEhrenbergrsquos Xanthidium ramosum as the type Spiniferitesramosus was redescribed by Davey amp Williams (1966) asbeing thin-walled with a smooth reticulate or granular wallwith gonal and intergonal processes They described sevenvarieties of this species with the typical variety Spiniferitesramosus var ramosus described as bearing gonal and occa-sional intergonal processes which are sometimes branchedThere was no description of the wall structure of the typevariety The size ranges reported by Davey amp Williams (1966)for Spiniferites ramosus var ramosus were 34ndash41 mm withprocess lengths 5ndash13 mm During the first workshop itbecame clear that most researchers use the description andillustrations of Rochon et al (1999 p 38 pl 9 figs 4ndash6) as aconceptual benchmark for Spiniferites ramosus In this senseSpiniferites ramosus has a smooth wall exclusively gonallong processes with low sutural crests and no apical bossthe reported central body length reported by Rochon et al(1999) is 30ndash46 mm LL suggested that this concept corre-sponds to Spiniferites ramosus subsp ramosus KNM said thatthe description of Spiniferites ramosus subsp ramosus ismuch broader and therefore it would be less ambiguous touse ldquoSpiniferites ramosus sensu Rochon et al 1999rdquo as donehere to refer to this particular morphology encountered inthe Quaternary This form is further discussed by Gurdebekeet al (2018)

KNM considered that Spiniferites ramosus var ramosussensu Davey amp Rogers 1975 cannot be attributed toSpiniferites ramosus sensu Rochon et al (1999) but shouldrather be attributed to Spiniferites belerius or Spiniferites bul-loideus sensu Wall 1965

MJH noted in draft that specimens described and illus-trated by Rochon et al (1999) were in fact referred toSpiniferites ramosus sensu lato and calling them ldquoSpiniferitesramosus sensu Rochon et al 1999rdquo does not fully capture theintent of these authors although understandably ldquoSpiniferitesramosus sensu lato sensu Rochon et al 1999rdquo is an impracti-cal label In the longer term a solution using formal nomen-clature would be preferable

231 Spiniferites ramosus subsp multiplicatus(Rossignol 1964) Lentin amp Williams 1973

Synonymy Hystrichosphaera furcata var multiplicataRossignol 1964 pl 1 fig 14 pl 3 fig 16

34 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

ldquoHystrichosphaera ramosa var multiplicatardquo Harland ampDownie 1969 fig 5 [combination not validly published] nonSpiniferites hyperacanthus (Deflandre amp Cookson) Sarjeant1970 [Matsuoka (1985a p 35) proposed the synonymy ofSpiniferites ramosus subsp multiplicatus with Spiniferiteshyperacanthus (Deflandre amp Cookson) Sarjeant 1970 but wefollow de Vernal et al (1992 p 325) and Londeix et al(2009 p 66) in not accepting this synonymy]Holotype Rossignol 1964 pl 1 fig 14 de Vernal et al 1992pl 8 fig 9Type locality Ashdod borehole coastal plain IsraelType stratum Pleistocene or Holocene (Rossignol 1964)Etymology Although not mentioned by Rossignol (1964)from Latin multiplicus (composed of many elements) pre-sumably in reference to the presence of intergo-nal processesDistinguishing characters Central body is ovoid bearingone or two intergonal processes that are bifurcated and pre-sent on every suture The wall is thick ndash comparable in thick-ness to Spiniferites pachydermus ndash with a smooth to granularwall surface The shape size and tabulation of the cyst aresimilar to those of Hystrichosphaera furcata (now considereda synonym of Spiniferites ramosus) The two main dorsalantapical processes are pronounced (Based on Rossignol1964 p 86)Dimensions Central body 44 x 40 mm process length15ndash20 mm (Rossignol 1964)Biological affinity UnknownIntraspecific morphotypes NoneComparison The presence of one or two occasional intergo-nal processes distinguishes this subspecies from Spiniferitesramosus subsp ramosus and from Spiniferites bulloideus Itdiffers from Spiniferites hyperacanthus by having only occa-sional intergonal processes by its more ovoid shape and thecommon occurrence of septaRemarks This taxon has been recorded only by Harland ampDownie (1969 pl 7 fig 5) Bradford amp Wall (1984 pl 4 figs15ndash17 pl 5 figs 1ndash3 5ndash7) and Londeix et al (2009 p 66not depicted) During the second workshop KNM wonderedif the specimens that he identified as Spiniferites ludhamensisfrom topotype material from the Ashdod borehole werewhat Rossignol (1964) had described and depicted asHystrichosphaera furcata var multiplicata KNM noted in draftthat Rossignol (1964 p 86) specified that ldquoThis variety mayhave a wall thickness similar to the variety pachydermardquohowever all other details of the wall structure are lacking LLexplained in draft that he considers Spiniferites ramosussubsp multiplicatus to accord with the concept illustrated bythe holotype drawing of Spiniferites ramosus subsp multipli-catus ndash that is to say similar to Spiniferites ramosus var ramo-sus but with consistent intergonal processes For him theconsistent presence of intergonal processes is the mostimportant chartacteristic for Spiniferites ramosus subspmultiplicatus as it was the first characteristic noted byRossignol in her original description of this taxon and is fea-tured in the name LL mentioned that the thick and granularwall that this subspecies ldquomay haverdquo is uncommon and

allows for variation in wall structure between smoothand granular

232 Spiniferites rhizophorus Head in Head amp Westphal1999 (Plate 6 figures 1ndash15)

Synonymy NoneHolotype Head amp Westphal 1999 fig 6 nos 1ndash4Type locality Clino Borehole Great Bahama BankCaribbean SeaType stratum Upper Lower Pliocene (Head ampWestphal 1999)Etymology Named in reference to the stilt-like branching ofprocess bases which recalls the aerial roots of the mangroveRhizophora (Head amp Westphal 1999)Distinguishing characters The central body wall appearsunstratified under light microscopy and has a nearly smooth(finely and faintly granulatepunctate) tegillum The centralbody is broadly ovoid with a very small apical protuberanceThe processes are gonal trifurcate (usually with secondarybifurcations) and solid with some branched and stilt-likebases reminiscent of the aerial roots of the mangrove treeRhizophora The sutures are reflected by faint lines or lowsolid ridges The apex is indicated by a tapering spine-likehollow process with smaller side branches The archeopyle isformed by loss of plate 3rdquo slightly reduced (Based on Headamp Westphal 1999 p 15ndash17)Dimensions Central body length 38 (460) 51 mm averageprocess length 9 (140) 17 mm (Head amp Westphal 1999)Biological affinity UnknownIntraspecific morphotypes NoneComparison During the first workshop MJH remarked thatthe drawings of the processes of Spiniferites tripodes inMorzadec-Kerfourn (1966 as Baltisphaeridium tripodes) aresimilar to the stilt-like columns of the processes of Spiniferitesrhizophorus and he suggested that the holotype or topotypematerial of the former species should be examined for com-parison with Spiniferites rhizophorus Presently the unclearoriginal description and illustration of Spiniferites tripodesdoes not permit a detailed comparisonRemarks None

233 Spiniferites ristingensis Head 2007 (Plate 7figures 1ndash15)

Synonymy Spiniferites sp 1 Head et al 2005 figs 9dndashgHolotype Head 2007 figs 8cndashlType locality Ristinge Klint DenmarkType stratum Eemian sediments of the Baltic Sea(Head 2007)Etymology Named after the type locality (Head 2007)Distinguishing characters This species has an ovoid centralbody with or without a short apical protuberance and bearsmembranous gonal processes joined by low sutural crestsSome processes are distally expanded to form irregular pol-ygonal platforms The central body wall is formed of twowall layers of similar thickness The pedium is smooth andthe tegillum forms small densely distributed blisters and

PALYNOLOGY 35

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

hollow undulations over the surface (losely comparable tobubble-wrap) The processes and sutural crests are formedby the tegillum and are distinctly bilayered The surface ofthe processes and sutural crests appears granulate (Basedon Head 2007 p 1011ndash1012)Dimensions Central body length 39 (430) 49mm wall thick-ness 10ndash18mm maximum process length 11 (129) 17 mm(Head 2007)Biological affinity UnknownIntraspecific morphotypes None knownComparison Spiniferites ristingensis is distinguished by itsmembranous processes and the distinctive wall structure of thecentral body Spiniferites ludhamensis from the Upper Plioceneof eastern England has a similar wall structure but has hollownon-membranous processes and hollow sutural crestsSpiniferites delicatus from modern sediments off the British Isleshas processes of similar shape but connected by high suturalcrests and a central-body wall structure characterised by a ped-ium with radial fibers and a thin granular tegillum whose sur-face appears microgranular to microreticulate Spiniferitesdelicatus also differs in having a reduced archeopyle (Based onHead 2007) For Spiniferites alaskensis see section 26Remarks During the first workshop VP wondered whetherthe wall structure of Spiniferites ristingensis is comparable tothe wall structure of Hafniasphaera MJH and LL did not thinkthis is comparable to Hafniasphaera because the wall struc-ture of Spiniferites ristingensis is not firmly vacuolate as inHafniasphaera During the second workshop there was alsodoubt as to whether the difference between Spiniferites ris-tingensis and Spiniferites delicatus warrants two separate spe-cies KNM mentioned during draft that he has seen thisspecies in recent sediments in the Gulf of Cadiz offshorePortugal the Baltic and Black seas and Baie de la Vilaine(Brittany) SR added during drafting that Spiniferites ristingen-sis is abundant (up to about 10 of the cyst assemblages) inmodern sediments off southwestern Portugal where itoccurs with Spiniferites delicatus

234 Spiniferites scabratus (Wall 1967) Sarjeant 1970(Plate 8 figures 1ndash6)

Synonymy Hystrichosphaera scabrata Wall 1967 p 102 pl14 figs 10ndash13 text-fig 2Holotype Wall 1967 pl 14 figs 10ndash13 Harland 1983 pl 45fig 7Type locality Core A254330 Yucatan Basin 19 35rsquo N 84

51rsquo W Caribbean SeaType stratum Holocene depth 150 cm (Wall 1967)Etymology Although not mentioned by Wall (1967) the epi-thet presumably refers to a scabrate tegillumDistinguishing characters The central body is ovoid with acingular displacement of one cingulum width The septa aremicrogranular and undulating The processes are exclusivelygonal there is a complex process at the anterior end of thesulcus There are three to four apical plates The archeopyleis rounded The sulcus is straight with the sulcal plates some-times visible particularly the posterior sulcal plate (Based onWall 1967 p 102)

Dimensions Central body length 48ndash55 mm process length10ndash17 mm (Wall 1967)Biological affinity The cystndashtheca relationship wasdescribed but not illustrated by Wall amp Dale (1968 p 271)as follows ldquoA resting spore identified as Hystrichosphaerascabrata Wall (using fossil terminology) from the Gulf ofParia was incubated to give a motile Gonyaulax theca of thespinifera type (48 x 38 mm) without antapical spines and withvery indistinct ornamentation The theca was very delicateand could not be identified with any speciesrdquoIntraspecific morphotypes A similar cyst was depicted byMatsuoka (2005) as Spiniferites sp cf scabratus in Holocenesediments off Santa Cruz Island (Galapagos) The differencebetween Matsuokarsquos form and Spiniferites scabratus isnot clearComparison Wall (1967) remarked that the differencebetween Spiniferites scabratus and Spiniferites membranaceusis the presence of a strong antapical flange in the latterRemarks MJH noted that it is not clear whether one or twospecimens were assigned as holotype and thus questionedthe validity of the name However personal communicationwith David Wall confirmed that it is a single specimen thatwas rotated into different views KZ remarked that the wall isnearly smooth and it is hard to see scabrate septa PJMstated that the holotype looks oxidised and the slide is inpoor shape KNM said that the processes are relatively broadand short and also noted that this species was recordedfrom Trondheimsfjord Norway by Dale (1976 pl 1 fig 6)However at the second workshop a specimen considered tobe Spiniferites scabratus by BD was identified as Spiniferitesristingensis by MJH and KNM MJH in draft provided the fol-lowing observations of the holotype The central body isovoidal with no apical protuberance The wall surfaceappears strongly granulate in plan view although uponcloser inspection the granules seem to be discrete solid rodsforming a columellate layer that separates two thin walllayers The rods are 06mm or less in diameter and are sep-arated from one another by their own width The entire wallthickness is 10 mm Processes have a scabrate surface arehollow and wide at their base and terminate in slender tri-furcations that project at right angles Only gonal processeswere seen Adjoining sutural crests are hollow The archeo-pyle is not reduced its margin closely following the sutures

235 Spiniferites septentrionalis Harland 1977 (Plate 8figures 7ndash9)

Synonymy Spiniferites aquilonius Matsuoka amp Bujak 1988p 74ndash76 pl 11 figs 6andashd pl 12 figs 1andashb pl 19 figs 4andashc7 text-figs 17AndashE [fide Matsuoka in Head amp Wrenn (1992 p26)] non Achomosphaera andalousiensis Jan du Chene 1977non Spiniferites ramuliferus (Deflandre 1937) Reid 1974Holotype Harland 1977 pl 1 figs 17ndash18Type locality Borehole SLN 7533 north-central North SeaType stratum Upper Quaternary (Harland 1977)Etymology From the Latin septentrionalis (northern north-erly) in reference to the occurrence of the type material inNorth Sea sediments (Harland 1977)

36 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Distinguishing characters Central body ovoid to spheroidalwith a shagreenate to scabrate or papillate outer wall withor without a small apical boss Generally no tabulation is vis-ible except for the archeopyle Processes are long slenderand gonal only The shafts of the more membranous proc-esses are commonly moderately perforate The processesvary distally from being trifurcate with bifid tips to being tri-furcate with perforate or fenestrate distal ends the latterstructure is especially prominent on the cingular processesThe archeopyle is simple reduced and formed by the loss ofplate 3rdquo (Based on Harland 1977 p 103ndash104 and newobservations of the holotype made by LL and KNM)Dimensions Central body length 3375 (4052) 4750mmwidth 2750 (3104) 3750mm wall thickness 10ndash20 mm pro-cess length 1000 (1240) 1625mm (Harland 1977)Biological affinity UnknownIntraspecific morphotypes NoneComparison Although Achomosphaera andalousiensis was con-sidered a synonym of Spiniferites septentrionalis by Harland(1983 p 326) and Jan du Chene amp Londeix (1988 p 421) thesynonymy was questioned by Head amp Wrenn (1992 p 2)During the first workshop participants expressed a general diffi-culty in understanding the difference between Achomosphaeraandalousiensis and Spiniferites septentrionalis and wondered ifthey could be synonymous During the first workshop LL con-sidered specimens illustrated by SEM in Harland (1988 pl 81as Achomosphaera andalousiensis) to belong to a species otherthan Achomosphaera andalousiensis (particularly specimen MPK5925) LL and MJH expressed the need to examine the holo-type of Spiniferites septentrionalis to determine whether or notit is conspecific with Achomosphaera andalousiensis In draftJames B Riding subsequently provided numerous photographsof the holotype Through study of these new images LL andKNM confirmed Spiniferites septentrionalis can be separatedfrom Achomosphaera andalousiensis on the basis of having amore elongate central body a shagreenate to scabrate or papil-late wall (sensu Williams et al 2000) and distal terminations ofthe processes that are fenestrate (rather than trabeculate) thefenestrations may not be on all trifurcations a contrast to themore regular process morphology in Achomosphaera andalou-siensis However MJH remarked in draft that the differencebetween the two species is not so straightforward and that aSEM micrograph of a specimen of Achomosphaera andalousien-sis in Harland 1988 (Plate 81 Figure 4) shows a transition tothe genus Spiniferites LL moreover suggested in draft that ldquoifyou look attentively at the central body surface of Spiniferitesseptentrionalis there are faint ridges and almost always septaaround plate 3rdquo In addition to the differences in process termi-nations Achomosphaera andalousiensis sometimes exhibits faintridges but no septa (see SEM micrographs in Jan du Chene1977 Jan du Chene amp Londeix 1988 Warny 1999) To keepSpiniferites septentrionalis in the genus Spiniferites is a way toretain this difference That is why I do not support transferringSpiniferites septentrionalis to Achomosphaerardquo Therefore thesynonymy between Achomosphaera andalousiensis andSpiniferites septentrionalis is not followed here See further dis-cussion in Londeix et al (2018) The perforate process bases are

similar to Spiniferites lazus but the latter does not possess fenes-trate distal ends (Harland 1977 p 103)Remarks Harland (1977 p 103) mentioned that Reid (perscomm to Harland) is of the opinion that Spiniferites ramulife-rus as recorded by him (1974) is synonymous withSpiniferites septentrionalis but we consider Reidrsquos specimento belong to Achomosphaera ramosasimilis

236 Spiniferites spinatus (Song Zhichen in SongZhichen et al 1985) Lentin and Williams 1989

Synonymy Spiniferites cingulatus (Wetzel 1933) Sarjeant 1970 varspinatus Song Zhichen in Song Zhichen et al 1985 p 43 pl 2 fig 5Holotype Song Zhichen in Song Zhichen et al 1985 pl 2fig 5Type locality 300 km east of Hangzhou Bay East China SeaType stratum Lower Quaternary (Song Zhichen et al 1985)Etymology Not provided by Song Zhichen in Song Zhichenet al (1985) but probably from the Latin spinate for spine-bearing in reference to the occurrence of processesDistinguishing characters Central body ovoid to subspher-oidal the epicyst and hypocyst being equal in size and sepa-rated by a wide and pronounced cingulum The cyst wallprobably consists of two layers and is relatively thin with acoarse to granular surface Sutures form smooth ridges2ndash3mm high At junctions there are short tubular processeswhich are narrow and hollow with trifurcate or bifurcate distalends Between each pair of gonal processes are at least twointergonal processes each with a wide base that abruptlytapers to a pointed end distally Tabulation is typical for thegenus but the archeopyle is not clear (Based on SongZhichen in Song Zhichen et al 1985 p 43 translation by HG)Dimensions Central body diameter 38 mm process length5 mm (Song Zhichen in Song Zhichen et al 1985)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks This taxon was initially described as a process-bearing variety of Spiniferites cingulatus (Wetzel 1933)Sarjeant 1970 (now Pterodinium cingulatum (Wetzel 1933)Below 1981) from the Senonian (Cretaceous) a species thatis probably not closely related It is difficult to assess whetherSpiniferites spinatus is conspecific with Spiniferites hyperacan-thus given the lack of a detailed description and an unclearimage We therefore suggested that the name Spiniferites spi-natus be restricted to the holotype

237 Spiniferites strictus Matsuoka 1983 (Plate 8figures 10ndash12)

Synonymy NoneHolotype Matsuoka 1983 pl 12 figs 5andashbType locality Takani-shinden Nishiyama-cho NiigataPrefecture central JapanType stratum Nishiyama Formation Pliocene or younger(Matsuoka 1983)Etymology From the Latin strictus (drawn tight) in referenceto the stout processes

PALYNOLOGY 37

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Distinguishing characters The central body is subspheroi-dal to ovoid with a smooth to granular outer wall somespecimens have a small apical boss The processes are bothgonal and intergonal and the process bases are sometimesperforated (Based on Matsuoka 1983 p 136ndash137)Dimensions Central body length 53ndash67 mm width50ndash62 mm process length 10ndash14 mm holotype wall thickness2mm (Matsuoka 1983)Biological affinity UnknownIntraspecific morphotypes NoneComparison Matsuoka (1983) stated that Spiniferites strictusis similar to Spiniferites bentorii but differs by having manyshort intergonal processesRemarks VP remarked during the first workshop thatSpiniferites strictus is similar to Spiniferites bentorii but notedthat she had not seen the material However LL did notagree that the two species look similar This species is dis-cussed by Limoges et al (2018)

238 Spiniferites tripodes (Morzadec-Kerfourn 1966)Lentin amp Williams 1973

Synonymy Baltisphaeridium tripodes Morzadec-Kerfourn1966 p 140ndash141 pl 3 figs 3ndash4Holotype Morzadec-Kerfourn 1966 pl 3 figs 3ndash4Type locality Borehole 36 drilled in 1961ndash1962 Marais dela Vilaine south of Redon FranceType stratum Holocene (Morzadec-Kerfourn 1966)Etymology From the Latin tri (three) and podes (feet)(Morzadec-Kerfourn 1966)Distinguishing characters The central body is spheroidaland smooth walled The base and top of each appendageare separate and some fenestrations may occur at mid-length Based on Morzadec-Kerfourn (1966 p 140ndash141)Dimensions Central body diameter 45mm average processlength 18 mm (Morzadec-Kerfourn 1966)Biological affinity UnknownMorphotypes NoneComparison See Spiniferites ristingensisRemarks Spiniferites tripodes was also recorded by Morzadec-Kerfourn (1979 from the Pelagian Sea east of Tunisia) and byMorzadec-Kerfourn (1984 from the Rhone Delta as Spiniferitescf tripodes) During the first workshop there was generalagreement that this species is not well understood LL notedthat it has been illustrated three times by Morzadec-Kerfournand each time it looked different We suggest that for nowSpiniferites tripodes be restricted to the holotype

3 Species previously assigned to Spiniferites buthere considered or accepted as belonging toother genera

31 Impagidinium inaequalis (Wall amp Dale in Wallet al 1973) Londeix et al 2009 (Plate 9figures 1ndash6)

Synonymy Spiniferites inaequalis Wall amp Dale in Wall et al1973 p 22 pl 1 figs 7ndash8

Holotype Wall et al 1973 pl 1 figs 7ndash8Type locality Core 1474P southeastern Black SeaType stratum Lower Holocene 725 cm (Wall et al 1973)Etymology Presumably in reference to the uneven shape ofthe broad hypocyst compared with the flask-like (lageni-form) epicystDistinguishing characters Cysts with a lageniform (flask-like) shape and an unevenly broad hypocyst The lageniformepicyst is subcylindrical to campanulate with weakly taperinglateral surfaces it has a smoothly rounded apex with anapical boss and expanded posterior precingular surfaces Thecingulum is situated at the cystrsquos mid-length and is in theform of a descending spiral the ends displaced by one cin-gulum width The ldquolopsidedrdquo hypocyst is broader than theepicyst and is irregularly subrectangular with a broad flatantapex the width of which equals that of the cyst near thecingulum The sulcus is almost vertical and narrow andequivalent in width to approximately one fifth of the widthof the hypocyst except at the antapex where it expands Inlateral view the cyst is irregularly subrectangular A perfecttabulation is reflected by low sutural ridges Several featuresof the tabulation are of special interest On the epicyst themid-ventral anterior sulcal plate is extensive and contactsboth 500 and 600 On the hypocyst plate 1p is large and sub-rectangular while the antapical plate 100 00 and the sulcalplates are transversely elongated Very small gonal spinesoccur on some specimens sometimes barely visible Thearcheopyle is formed by loss of plate 3rdquo The wall is scabrateto microgranular (Based on Wall et al 1973 p 22)Dimensions Central body length (without processes)44ndash55 mm width 33ndash43 mm processes up to 5 mm long (Wallet al 1973)Biological affinity UnknownIntraspecific morphotypes NoneComparison The peculiar shape of the central body makesthis species uniqueRemarks The species was also found in recent sedimentsfrom the Black Sea and Marmara Sea by Mudie et al (2002)During the first workshop LL explained that he had reattrib-uted Spiniferites inaequalis to Impagidinium because occa-sional ldquovery small gonal spinesrdquo or ldquominute projectionsrdquo isnot enough to assign the species to the genus SpiniferitesSpiniferites cingulatus and Spiniferites crassimuratus (Davey ampWilliams 1966) Sarjeant 1970 were transferred for similar rea-sons to Pterodinium by Below (1981) and Thurow et al(1988) respectively Although not specified by Londeix et al(2009) the species also accords with Impagidinium becauseplate 6 is triangular and contacts 4 (Wall et al 1973 pl 1fig 8) PJM noted that she has observed well-preservedspecimens of this species in mid-Pleistocene to Holocenesediments of the northern Caspian Sea (Volga and Emba del-tas) as well as in Upper Miocene material from the Black Seaprovided by FS BD stated in draft that ldquovery small spines orminute projections are accepted for maintaining species asecophenotypes within the genera OperculodiniumLingulodinium and Spiniferites from low salinity environmentssuch as the Black Sea and the Baltic (Dale 1996) and there-fore he accepts Spiniferites inaequalis within Spiniferites MJH

38 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

added in draft his following observations of the holotype Itscentral body has a slight apical protuberance about 15mmhigh a fairly thick (08ndash10mm) wall and an irregularlygranulo-reticulate surface with murigranules about 03mmor less in width The sutural crests reach 20mm high andare solid except at the base (minutely suturocavate)Processes are gonal up to 10 mm in length solidunbranched and taper to blunt points The cingulum dis-placed by about 1 width The archeopyle margin follows thesutures quite closely MJH further remarked that it may bepremature to transfer this species to the genusImpagidinium The processes while small for a species ofSpiniferites are discrete elements rather than extensions ofthe crests as found for example in I aculeatum Given theextreme morphological plasticity of many Paratethyan formsMJH wonders how Spiniferites inaequalis compares with thefull range of morphology exhibited by Spiniferites cruciformiswhich is found in the same samples He presently acceptsthe name Spiniferites inaequalis

32 Spiniferites rubinus (Rossignol 1962 ex Rossignol1964) Sarjeant 1970

Synonymy ldquoHystrichosphaeridium rubinardquo Rossignol 1962 p134 [invalid because no illustration provided]

Hystrichosphaera rubina Rossignol 1962 p 134 exRossignol 1964 p 87ndash88 pl 1 figs 12ndash13 pl 3 figs 22ndash23Holotype Rossignol 1964 pl 1 figs 12ndash13Type locality Borehole close to Rubin 230 coastalplain IsraelType stratum Quaternary depth 160ndash166 m(Rossignol 1964)Etymology Named after a stream called Wadi Rubin(Rossignol 1964)Distinguishing characters An ovoid cyst with a micropunc-tate or faintly and finely granular wall that also makes upthe processes and sutural membranes The tabulation isunclear but plate areas 4and 1 can be recognised The cin-gulum and sulcus are also usually conspicuous but do notexhibit individual plates The sulcus is broad and the cingu-lum is a laevorotatory helicoid planar to displaced by one-half of its width and not indented Processes are gonal andmembranous and are most conspicuous in the apical antapi-cal cingular and sulcal areas They are somewhat flexuousand have petaloid tips Some specimens have many mem-branes especially in the areas listed The sulcus is so broadand is often surrounded by such high membranes that thecyst appears to bear membranes only around its marginHowever membranes may not be developed between thesulcus and the adjacent pre- and postcingular regions Inthese forms some of the non-membranous processes appearas small protuberances Complex boxlike processes alsooccur at the apex of the sulcus and in the position of theposterior intercalary plate Some of the more complex mem-branous processes carry small spines In specimens thatexhibit more obvious processes those in the cingular regionare erect simple slender and conical The archeopyle is

formed by the loss of plate 3rdquo (Based on Rossignol 1964 p134 Harland 1979 p 537 and Head 1996b p 560)Dimensions Central body length 52 mm height 44 mm crestheight 15ndash20mm (Rossignol 1964) Central body length 46(530) 61 mm equatorial diameter 47 (495) 53 mm maximumcrest height 10ndash11mm (Head 1996b) Central body lengthexcluding processes or membranes 460ndash540 lm width380ndash460lm maximum process length or membrane height100ndash120lm (Harland 1979)Biological affinity UnknownIntraspecific morphotypes NoneComparison NoneRemarks During the first workshop KNM questionedwhether this species really is a Spiniferites MJH remarkedthat it is like a Spiniferites but also resembles Invertocystalacrymosa Edwards 1984 LL asked ldquowhy not PentadiniumGerlach 1961rdquo MJH replied that when the tegillum retractsyou get the resemblance of a continuous spine-like structuresee for instance specimens illustrated by Harland (1979 pl 2fig 4ndash11) which represent one end of the spectrum LL con-sidered that it does not belong in Spiniferites but was uncer-tain where else it could go Since there are no processes onthis species it cannot remain in Spiniferites It is approachingImpagidinium (if the septa are solid) or Pentadinium (if thesepta form sutural pericoels) but could also be questionablyassigned to Invertocysta MJH agreed PJM remarked after thefirst workshop that ldquothe specimens of Spiniferites rubinus Ihave seen in the North Atlantic are clearly different fromInvertocysta in periphragmal attachment size and arrange-ment I have seen no signs of intermediatesrdquo There was con-sensus that this species does not belong to Spiniferites butthat no existing genus could be identified that would accom-modate it so for now the species is retained questionably inSpiniferites

33 Thalassiphora balcanica Baltes 1971

Synonymy Disphaeria balcanica (Baltes 1971 p 6 pl 3 figs3ndash7) Norvick 1976 p 99

ldquoSubathua balcanicardquo (Baltes 1971 p 6 pl 3 figs 3ndash7)Khanna amp Singh 1980 p 308 and 1981b p 394[Combination not validly published]

Spiniferites balcanicus (Baltes 1971 pl 3 figs 3ndash7) Seurouto-Szentai 2000 p 162Holotype Baltes 1971 pl 3 figs 3ndash7Type locality Pannonian Basin RomaniaType stratum Lower Pliocene Pontian regional stage(Baltes 1971)Etymology Presumably in reference to the Balkan Peninsulawhich includes RomaniaDistinguishing characters The description of Baltes (1971p 6) indicates that the holotype of Thalassiphora balcanicafrom Romania is a very large (120 130mm) cyst with anellipsoid central body partly enfolded in a membranousouter wall that contacts the central body only on one side(dorsal side [sic]) Baltes described the wall surface as pter-ate and fibrous quasi-reticulate and perforate resembling alamellate wing Baltes described the central body as having a

PALYNOLOGY 39

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

vaguely outlined cingulum and a trapezoidal cingular arche-opyle but otherwise lacking tabulation However Seurouto-Szentai (2000 p 162) transferred Thalassiphora balcanica toSpiniferites as S balcanicus based on specimens from UpperPannonian deposits in boreholes from Hungary some ofwhich are illustrated as having highly reduced undecipher-able ornamentation and others with fenestrate luxuria andattachments like those of Galeacysta etrusca (see Mudieet al 2018) The emended diagnosis of Seurouto-Szentai (2000)for Spiniferites balcanicus describes an ovoid or spheroidalcentral body with two membranes ldquofixed both on the rightand left side of the sulcusrdquo and sometimes with two wing-like membranes that are ldquodifferently perforatedrdquo one ofthem being arched in the apical area and being partiallyconnected except in the ventral areaDimensions Central body length 90ndash100mm total length120ndash130 mm archeopyle 35ndash45 mm (Baltes 1971) Centralbody length 80ndash90 mm total length 80ndash110 mm (Seurouto-Szentai 2000)Biological affinity UnknownIntraspecific morphotypes NoneComparison At the second workshop PJM suggested thatthis taxon may be the same as the Late Eocene informal spe-cies ldquoThalassiphora subreticulatardquo of Fensome amp Williams(2005) The camocavate species described by Baltes clearlydiffers from Galeacysta etrusca which has a completely differ-ent non-fibrous tegillum and has a galeate (helmet-shaped)tegillum with tabulate claustra (ie very large arch-shapedcamerate fenestrations) and with a completely differentattachment to the endocyst than the membranous envelop-ing perforate tegillum of Thalassiphora balcanica In additionthe light microscope images of three specimens ofSpiniferites balcanicus from the Paks 4 borehole (Seurouto-Szentai2000 pl IX figs 1ndash3) neither demonstrate the features MariaSeurouto-Szentai describes nor do they support her differentialdiagnosis in which she distinguishes Spiniferites balcanicusfrom Subathua balcanicaRemarks Subathua balcanica is invalidly published becausethe basionym was not fully referenced Stover amp Evitt (1978p 194) considered Thalassiphora pelagica to be the seniorsynonym of this taxon

During the first workshop there was a general discussionas to why Thalassiphora balcanica might belong toSpiniferites as suggested by Seurouto-Szentai (2000) PJM addedin draft that this assignment was based on the comments inthe paper by Seurouto-Szentai (2000) that mention gonyaulacoidtabulation marked by ldquostumps of processesrdquo in some but notall cysts which ldquohellipprobably depends on stage of ontogenyrdquoHowever these processes have not been observed in sam-ples examined during the workshops and the cysts PJM hasseen show many characteristics of Thalassiphora Given theseobservations it seems prudent to assume that the speciesindeed belongs to Thalassiphora since it had been originallydescribed as Thalassiphora baltica The suggestion was alsoexpressed that the species could belong toHystrichostrogylon There was general agreement that this isnot a Spiniferites species and therefore we retain it inThalassiphora

Mudie et al (2018) found no intergradation in morph-ology between Thalassiphora balcanica and Galaecystaetrusca other than possibly some overlap at the upper endof their endocyst-ectocyst size and these authors agree toretain Thalassiphora balcanica PJM stated that the locationof the holotype (Baltesrsquos specimen LC 6229 -44106) is nowunknown and the lectotype designated by Seurouto-Szentai(2000) as Spiniferites balcanicus held by Maria Seurouto-Szentai atKomlo in southwestern Hungary are severely dessicatedrequiring new studies of material from the type localityDetailed morphological studies of specimens from northernCroatia are currently under investigation by PJM AR RobFensome and Koraljke Bakrac (Croatian Geological Survey)

4 Discussion about genusspeciesvariety concepts

During the round table discussion held at the second work-shop several themes were discussed the major foci being 1)what to do with problematic species 2) concepts inofSpiniferites taxonomy (what constitutes a genus species var-iety and forma) and 3) dual versus unified taxonomy (seealso Ellegaard et al 2018) This discussion can be found inSupplementary Appendix 3

Acknowledgements

KNM and MJH extend their appreciation to David Wall for sending holo-type and topotype material and James B Riding for rephotographingthe holotype of Spiniferites septentrionalis Rob Fensome is thanked forhis constructive and insightful review

References

Baltes N 1971 Pliocene Dinoflagellata and Acritarcha in Romania InFarinacci A editor 2nd Planktonic Conference Rome 1970Proceedings Rome Italy Edizioni Tecnoscienza p 1ndash16

Below R 1981 Dinoflagellaten-Zysten aus dem oberen Hauterive bisunteren Cenoman Seuroud-West-Marokkos Palaeontogr Abt B 1761ndash145

Below R 1982 Scolochorate Zysten der Gonyaulacaceae (Dinophyceae)aus der Unterkreide Marokkos Palaeontogr Abt B 1821ndash51

Bradford MR Wall DA 1984 The distribution of Recent organic-walleddinoflagellate cysts in the Persian Gulf Gulf of Oman and northwest-ern Arabian Sea Palaeontogr Abt B 19216ndash84

Bujak JP 1984 Cenozoic dinoflagellate cysts and acritarchs from theBering Sea and northern North Pacific DSDP Leg 19Micropaleontology 30(2)180ndash212

Bujak JP Matsuoka K 1986 Taxonomic reallocation of Cenozoic dinofla-gellate cysts from Japan and the Bering Sea Palynology 10(1)235ndash241

Beuroutschli O 1885 II Abtheilung Mastigophora In Dr HG BronnrsquosKlassen und Ordnungen des Thier-Reichs wissenschaftlich dargestelltin Wort und Bild Ester Band Protozoa Leipzig amp Heidelberg CFWinterrsquosche Verlagsbuchhandlung p 865ndash1088

Claparede E Lachmann J 1859 Etudes sur les infusoires et les rhizo-podes Institut National Genevois Memoires 6(Memoire 1)261ndash482[Cover date 1858 issue date 1859 fide Fensome et al 1993 p 218]

Cookson IC Eisenack A 1974 Mikroplankton aus australischen mesozoi-schen und tertieuroaren Sedimenten Palaeontogr Abt B 14844ndash93

Corradini D Biffi U 1988 Etude des dinokystes a la limite Messinien-Pliocene dans la coupe Cava Serredi Toscane Italie Dinocyst study atthe Messinian-Pliocene boundary in the Cava Serredi sectionTuscany Italy Bulletin Des Centres de Recherches Exploration-Production Elf-Aquitaine 12221ndash236

40 K N MERTENS ET AL

Dale B 1976 Cyst formation sedimentation and preservation factorsaffecting dinoflagellate assemblages in Recent sediments fromTrondheimsfjord Norway Rev Palaeobot Palynol 22(1)39ndash60

Dale B 1983 Dinoflagellate resting cysts benthic plankton In FryxellGA editor Survival strategies of the algae Cambridge CambridgeUniversity Press p 69ndash136

Dale B 1996 Dinoflagellate cyst ecology modelling and geologicalapplications In Jansonius J McGregor DC editors PalynologyPrinciples and Applications volume 3 AASP Foundation Dallas TX p1249ndash1275

Dale B Dale AL Prince I 2005 Statistical modelling of ecological signalsa new method for biostratigraphy In Powell AJ Riding JB editorsRecent Developments in Applied Biostratigraphy London (UK) TheMicropalaeontological Society Special Publications p 179ndash203

Davey RJ Rogers J 1975 Palynomorph distribution in recent offshoresediments along two traverses off South West Africa Mar Geol 18(4)213ndash225

Davey RJ Williams GL 1966 IV The genera Hystrichosphaera andAchomosphaera In Davey RJ Downie C Sarjeant WAS Williams GLeditors Studies on Mesozoic and Cainozoic dinoflagellate cysts BritishMuseum (Natural History) Geology Bulletin Supplement 3 p 28ndash52

Deflandre G 1937 Microfossiles des Silex Cretaces Deuxieme PartieFlagelles incertae sedis Hystrichosphaeridees - SarcodinesOrganismes divers Annales Paleont 2651ndash103

Deflandre G Cookson IC 1955 Fossil microplankton from Australian LateMesozoic and Tertiary sediments Mar Freshwater Res 6(2)242ndash313

Demetrescu E 1989 Achomosphaera argesensis a new dinoflagellatespecies from the early Pliocene of the southern Carpathians foredeepRomania Rev Palaeobot Palynol 59(1-4)51ndash55

De Schepper S Head MJ Louwye S 2004 New dinoflagellate cyst andincertae sedis taxa from the Pliocene of northern Belgium southernNorth Sea basin J Paleo 78(4)625ndash644

de Vernal A Londeix L Mudie PJ Harland R Morzadec-Kerfourn MTuron J-L Wrenn JH 1992 Quaternary organic-walled dinoflagellatecysts of the North Atlantic Ocean and adjacent seas ecostratigraphyand biostratigraphy In Head MJ Wrenn JH editors Neogene andQuaternary Dinoflagellate Cysts and Acritarchs Dallas USAAmerican Association of Stratigraphic Palynologists Foundation p289ndash328

de Vernal A Mudie PJ 1992 Pliocene and Quaternary dinoflagellate cyststratigraphy in Labrador Sea paleoecological implications Neogeneand Quaternary dinoflagellate cysts and acritarchs In Head MJWrenn JH editors College Station (TX) American Association ofStratigraphic Palynologists Foundation Houston TX p 329ndash46

Diesing KM 1866 Revision der Prothelminthen AbtheilungMastigophoren Akademie Der Wissenschaften zu WienSitzungsberichte Mathematisch-Naturwissenschaftlige Klasse 52287ndash401

Dodge JD 1985 Atlas of Dinoflagellates London Farrand PressDodge JD 1989 Some revisions of the family Gonyaulacaceae

(Dinophyceae) based on scanning electron microscope study BotMar 32275ndash298

Dybkjaer K Piasecki S 2010 Neogene dinocyst zonation for the easternNorth Sea Basin Denmark Rev Palaeobot Palynol 161(1-2)1ndash29

Eaton GL 1996 Seriliodinium a new Late Cenozoic dinoflagellate fromthe Black Sea Rev Palaeobot Palynol 91(1-4)151ndash169

Edwards LE 1984 Miocene dinocysts from Deep Sea Drilling Project Leg81 Rockall Plateau eastern North Atlantic Ocean In Roberts DGSchnitker D editors Deep Sea Drilling Project Washington InitialReports 81 Washington DC Government Printing Office p 581ndash594

Ehrenberg CG 1837 euroUber das Massenverheuroaltnis der jetzt lebendenKiesel-Infusorien und eurouber ein neues Infusorien-Conglomerat alsPolirschiefer von Jastraba in Ungarn Abh Akad Wiss Berlin (1836) 1109ndash135

Eisenack A 1954 Mikrofossilien aus Phosphoriten des samleuroandischenUnteroligozeuroans und eurouber die Einheitlichkeit der HystrichosphaerideenPalaeontogr Abt A 10549ndash95

Eisenack A Gocht H 1960 Neue Namen feurour einige Hystrichospheuroaren derBernsteinformation Ostpreussens Neues Jahrb Geol P M 11511ndash518

Ellegaard M Daugbjerg N Rochon A Lewis J Harding I 2003Morphological and LSU rDNA sequence variation within theGonyaulax spinifera-Spiniferites group (Dinophyceae) and proposal ofG elongata comb nov and G membranacea comb nov Phycologia42(2)151ndash164

Ellegaard M Head MJ Versteegh GJM 2018 Linking biological and geo-logical data on dinoflagellates using the genus Spiniferites as anexample the implications of species concepts taxonomy and dualnomenclature Palynology 42(Suppl1) httpdoiorg1010800191612220181465732

Ellegaard M Lewis J Harding I 2002 Gonyaulax baltica sp nov(Dinophyceae) - cyst-theca relationship life cycle and environmentallyinduced morphological variations in the cyst of a new species fromthe Baltic J Phycol 38(4)775ndash789

Evitt WR 1963 A discussion and proposals concerning fossil dinoflagel-lates hystrichospheres and acritarchs I National Academy ofSciences Washington Proceedings 49(2)158ndash164

Fensome RA Williams GL 2004 The Lentin and Williams Index of fossildinoflagellates 2004 Edition American Association of StratigraphicPalynologists Contributions Series 42909 p

Fensome RA Williams GL 2005 Scotian Margin PalyAtlas version 1Geological Survey of Canada Open File 677

Fensome RA Gocht H Stover LE Williams GL 1991 The Eisenack Catalogof Fossil Dinoflagellates New Series Volume 1 Stuttgart Germany ESchweizerbartrsquosche Verlagsbuchhandlung Stuttgart Germany

Fensome RA Taylor FJR Norris G Sarjeant WAS Wharton DI WilliamsGL 1993 A classification of fossil and living dinoflagellatesMicropaleontology Special Publication 71ndash245

Fuchs R Seurouto-Szentai M 1991 Organisches Mikroplankton(Phytoplankton) aus dem Pannonien des Wiener Beckens (euroOsterreich)und Korrelationsmeurooglichkeiten mit dem zentralen pannonischenBecken (Ungarn) Jubileuroaumsschrift 20 Jahre GeologischeZusammenarbeit euroOsterreich - Ungarn 119ndash34

Gibbard PL Head MJ 2010 The newly-ratified definition of theQuaternary SystemPeriod and redefinition of the Pleistocene SeriesEpoch and comparison of proposals advanced prior to formal ratifica-tion Episodes 33152ndash158

Gocht H 1969 Formengemeinschaften alttertieuroaren Mikroplanktons ausBohrproben des Erdeuroolfeldes Meckelfeld bei Hamburg PalaeontogrAbt B 1261ndash100

Gurdebeke P Mertens KN Bogus K Marret F Chomerat N Vrielinck HLouwye S 2018 Taxonomic re-investigation and geochemical charac-terization of Reidrsquos (1974) species of Spiniferites from holotype andtopotype material Palynology 42(Suppl1) httpdoiorg1010800191612220181465735

Hansen JM 1977 Dinoflagellate stratigraphy and echinoid distribution inUpper Maastrichtian and Danian deposits from Denmark Bulletin ofthe Geological Society of Denmark 261ndash26

Harland R 1968 A microplankton assemblage from the post-Pleistoceneof Wales Grana Palynologica 8(2-3)536ndash554

Harland R 1977 Recent and Late Quaternary (Flandrian and Devensian)dinoflagellate cysts from marine continental shelf sediments aroundthe British Isles Palaeontogr Abt B 16487ndash126

Harland R 1979 Dinoflagellate biostratigraphy of Neogene andQuaternary sediments at holes 400400A in the Bay of Biscay (DeepSea Drilling Project Leg 48) In Montadert L editors Deep SeaDrilling Project Washington Initial Reports 48 Washington DCGovernment printing office p 531ndash545

Harland R 1988 Quaternary dinoflagellate cyst biostratigraphy of theNorth Sea Palaeontology 31(3)877ndash903

Harland R Reid PC Dobell P Norris G 1980 Recent and sub-Recentdinoflagellate cysts from the Beaufort Sea Canadian Arctic Grana19(3)211ndash225

Harland R 1983 Distribution maps of Recent dinoflagellate cysts in bot-tom sediments from the North Atlantic Ocean and adjacent seasPalaeontology 26321ndash387

Duane A Harland R 1990 Quaternary dinoflagellate cyst biostratigraphyof the North Sea Palaeontology 63(1-2)1ndash903

Harland R Downie C 1969 The dinoflagellates of the interglacial depos-its at Kirmington Lincolnshire P Yorks Geol Soc 37(2)231ndash237

PALYNOLOGY 41

Harland R Sharp J 1986 Elongate Spiniferites cysts from North Atlanticbottom sediments Palynology 10(1)25ndash34

Head MJ 1994 Morphology and paleoenvironmental significance of theCenozoic dinoflagellate genera Tectatodinium and HabibacystaMicropaleontology 40(4)289ndash321

Head MJ 1996a Modern dinoflagellate cysts and their biological affin-ities In Jansonius J McGregor DC editors Palynology principles andapplications vol 3 Dallas USA American Association ofStratigraphic Palynologists Foundation p 1197ndash1248

Head MJ 1996b Late Cenozoic dinoflagellates from the Royal Societyborehole at Ludham Norfolk eastern England J Paleontol 70(04)543ndash570

Head MJ 1997 Thermophilic dinoflagellate assemblages from the midPliocene of eastern England J Paleontol 71(2)165ndash193

Head MJ 2007 Last Interglacial (Eemian) hydrographic conditions in thesouthwestern Baltic Sea based on dinoflagellate cysts from RistingeKlint Denmark Geol Mag 144(6)987ndash1013

Head MJ Gibbard PL 2015 Formal subdivision of the QuaternarySystemPeriod Past present and future Quaternary International3834ndash35

Head MJ Westphal H 1999 Palynology and paleoenvironments of aPliocene carbonate platform the Clino core Bahamas J Paleontol73(01)1ndash25

Head MJ Wrenn JH 1992 A forum on Neogene and Quaternary dinofla-gellate cysts In Head MJ Wrenn JH editors Neogene andQuaternary Dinoflagellate Cysts and Acritarchs Dallas USAAmerican Association of Stratigraphic Palynologists Foundation p1ndash31

Head MJ Seidenkrantz M-S Janczyk-Kopikowa Z Marks L Gibbard PL2005 Last Interglacial (Eemian) hydrographic conditions in the south-eastern Baltic Sea NE Europe based on dinoflagellate cysts Quat Int130(1)3ndash30

Head MJ Fensome RA Herendeen PS Skog JE 2016 (315ndash319)Proposals to amend Article 118 and its Examples to remove ambigu-ity in the sanctioning of dual nomenclature for dinoflagellates andan emendation of Article 117 Example 29 Taxon 65(4)902ndash903

Hyesu Y 1981 Dinoflagellaten aus der Oberkreide (Santon) vonWestfalen Palaeontogr Abt B 1771ndash89

Islam MA 1983 Dinoflagellate cysts from the Eocene of the London andthe Hampshire basins southern England Palynology 7(1)71ndash92

Jan Du Chene R 1977 Etude palynologique du Miocene superieurAndalou (Espagne) Rev Esp Micropaleontol 997ndash114

Jan Du Chene R Londeix L 1988 Donnees nouvelles sur Achomosphaeraandalousiense Jan du Chene 1977 kyste de dinoflagelle fossileBulletin Des Centres de Recherches ExplorationndashProductionElfndashAquitaine 12237ndash250

Khanna AK Singh HP 1980 Subathua - a new dinoflagellate genus andits palaeoecological significance in the Subathu Formation SimlaHills The Palaeobotanist 26307ndash313

Klumpp B 1953 Beitrag zur Kenntnis der Mikrofossilien des mittlerenund oberen Eozeuroan Palaeontogr Abt A 103377ndash406

Kofoid CA 1911 Dinoflagellata of the San Diego region IV The genusGonyaulax with notes on its skeletal morphology and a discussion ofits generic and specific characters University of California Publicationsin Zoology 8187ndash286

Lentin JK Williams GL 1973 Fossil dinoflagellates index to genera andspecies Geological Survey of Canada Paper no 73 42 176 p

Lentin JK Williams GL 1981 Fossil dinoflagellates index to genera andspecies 1981 edition Bedford Institute of Oceanography ReportSeries No BI-R-81ndash12 345 p

Lentin JK Williams GL 1989 Fossil dinoflagellates index to genera andspecies 1989 edition American Association of StratigraphicPalynologists Contributions Series 20473 p

Lewis J Rochon A Harding I 1999 Preliminary observations of cyst-theca relationships in Spiniferites ramosus and Spiniferites membrana-ceus (Dinophyceae) Grana Suppl 31ndash12

Limoges A Londeix L Mertens KN Rochon A Pospelova V Cuellar T deVernal A 2018 Identification key for Pliocene and QuaternarySpiniferites taxa bearing intergonal processes based on observations

from estuarine and coastal environments Palynology 42(Suppl1)httpdoiorg1010800191612220181465733

Londeix L Benzakour M de Vernal A Turon J-L Suc J-P 1999 LateNeogene dinoflagellate cyst assemblages from the Strait of Sicily cen-tral Mediterranean Sea paleoecological and biostratigraphical implica-tions In Wrenn JH Suc J-P Leroy SAG editors The Pliocene Time ofChange Dallas USA American Association of StratigraphicPalynologists Foundation p 65ndash91

Londeix L Herreyre Y Turon J-L Fletcher W 2009 Last Glacial toHolocene hydrology of the Marmara Sea inferred from a dinoflagel-late cyst record Rev Palaeobot Palynol 158(1-2)52ndash71

Londeix L Zonneveld K Masure E 2018 Taxonomy and operationalidentification of Quaternary species of Spiniferites and related generaPalynology 42(Suppl1) httpdoiorg1010800191612220181465740

Louwye S Head MJ De Schepper S 2004 Dinoflagellate cyst stratig-raphy and palaeoecology of the Pliocene in northern Belgium south-ern North Sea Basin Geol Mag 141(3)353ndash378

Mantell GA 1854 The Medals of Creation or First lessons in Geologyand the study of Organic Remains 2nd edn 2 vols London HGBohn

Marret F de Vernal A Pedersen TF McDonald D 2001 MiddlePleistocene to Holocene palynostratigraphy of Ocean Drilling ProgramSite 887 in the Gulf of Alaska northeastern North Pacific Can J EarthSci 38(3)373ndash386

Marret F Leroy S Chalie F Francoise F 2004 New organic-walled dino-flagellate cysts from recent sediments of central Asian seas RevPalaeobot Palynol 129(1-2)1ndash20

Marret F Mertens KN 2018 Additional observations of Spiniferites alas-kensis from topotype material Palynology 42(Suppl1) httpdoiorg1010800191612220181465734

Matsuoka K 1976 Paleoenvironmental study of the Saho and theSaidaiji Formations from a view point of palynology Mizunami FossilMuseum Bulletin 399ndash117

Matsuoka K 1983 Late Cenozoic dinoflagellates and acritarchs in theNiigata district central Japan Palaeontogr Abt B 18789ndash154

Matsuoka K 1985 Organic-walled dinoflagellate cysts from surface sedi-ments of Nagasaki Bay and Senzaki Bay west Japan Faculty ofLiberal Arts Nagasaki University Natural Science Bulletin 2521ndash115

Matsuoka K 1987a Organic-walled dinoflagellate cysts from surface sedi-ments of Akkeshi Bay and Lake Saroma North Japan Faculty ofLiberal Arts Nagasaki University Natural Science Bulletin 2835ndash123

Matsuoka K 1987b Investigation on fossil dinoflagellates - Dinoflagellatecyst assemblages of Tama Lowland (Loc 3) In Matsushita Y editorIntegrated studies on the Holocene sediment of Kawasaki City Japanp 83ndash88 [In Japanese]

Matsuoka K 1992 Species diversity of modern dinoflagellate cysts in sur-face sediments around the Japanese Islands In Head MJ Wrenn JHeditors Neogene and Quaternary Dinoflagellate Cysts and AcritarchsDallas USA Houston (TX) American Association of StratigraphicPalynologists Foundation pp 33ndash53

Matsuoka K 2005 Modern dinoflagellate cysts found in surface sedi-ments of Santa Cruz Island Galapagos Galapagos Research 638ndash11

Matsuoka K Bujak JP 1988 Cenozoic dinoflagellate cysts from theNavarin Basin Norton Sound and St George Basin Bering SeaNagasaki University Faculty of Liberal Arts Natural Science Bulletin29(1)1ndash147

Matsuoka K Fukuyo Y Jaafar MH de Silva MWRN 1989 Occurrence ofthe cyst of Pyrodinium bahamense var compressum in surface sedi-ments of Brunei Bay In Hallegraeff GM Maclean JL editors Biologyepidemiology and management of Pyrodinium red tides InternationalCenter for Living Aquatic Resources Management (Manila Philippines)Conference Proceedings 2189ndash95

McMinn A 1991 Recent dinoflagellate cysts from estuaries on thecentral coast of New South Wales Australia Micropaleontology 37(3)269ndash287

Mertens KN Aydin H Uzar S Takano Y Yamaguchi A Matsuoka K 2015Cyst-theca relationship and phylogenetic position of the dinoflagel-late cyst Spiniferites pachydermus from Izmir bay Turkey J Phycol51(3)560ndash573

42 K N MERTENS ET AL

Mertens KN Carbonell-Moore C 2018 Introduction to SpiniferitesMantell 1850 special issue Palynology 42(Suppl1) httpdoiorg1010800191612220181465741

Morquecho L Gongora-Gonzalez DT Okolodkov YB 2009 Cyst-thecarelationships of Gonyaulacales and Peridiniales (Dinophyceae) fromBahıa Concepcion Gulf of California Acta Bot Mex 88(88)9ndash29

Morzadec-Kerfourn M-T 1966 Etude des acritarches et dinoflagelles dessediments vaseux de la Vallee de la Vilaine aux environs de Redon(Ille-et-Vilaine) Bulletin de la Societe geologique et mineralogique deBretagne Nouvelle Serie 137ndash146

Morzadec-Kerfourn M-T 1979 D - Les kystes de Dinoflagelles InGeologie Mediterraneenne VI (1) La Mer Pelagienne Etudesedimentologique et ecologique du Plateau tunisien et du GolfedeGabes Editions de lrsquouniversite de Provence p 221ndash246

Morzadec-Kerfourn M-T 1984 Les kystes des dinoflagelles dans les sedi-ments Plehistocenes superieurs et Holocenes au large du delta duRhone et de la Corse Ecologie Des Microorganismes en MediterraneeOccidentale ECOMED Petroles et Techniques 170ndash183

Mudie PJ Aksu AE Yasar D 2001 Late Quaternary dinoflagellatecysts from the Black Marmara and Aegean seas variations in assemb-lages morphology and paleosalinity Mar Micropaleontol 43(1-2)155ndash178

Mudie PJ Rochon A Aksu AE Gillespie H 2002 Dinoflagellate cystsfreshwater algae and fungal spores as salinity indicators in LateQuaternary cores from Marmara and Black Seas Marine Geology 190203ndash231

Mudie p Rochon A Richards K Ferguson S Warny S 2018 Spiniferitescruciformis Pterocysta cruciformis and Galeacysta etrusca morph-ology and palaeoecology Palynology 42(Suppl1) httpdoiorg1010800191612220181465737

Norvick MS 1976 Mid-Cretaceous microplankton from Bathurst IslandIn Norvick MS Burger D editors Palynology of the Cenomanian ofBathurst Island Northern Territory Australia Bureau of MineralResources Geology and Geophysics Bulletin 15121ndash113

Popescu S-M Dalesme F Jouannic G Escarguel G Head MJ Melinte-Dobrinescu MC Seurouto-Szentai M Bakrac K Clauzon G Suc J-P 2009Galeacysta etrusca complex dinoflagellate cyst marker ofParatethyan influxes into the Mediterranean Sea before and afterthe peak of the Messinian Salinity Crisis Palynology 33(2)105ndash134

Pouchet G 1883 Contribution a lrsquoetude des cilioflagelles Journ AnatPhysiol Paris 19399ndash455

Price AM Pospelova V 2014 Spiniferites multisphaerus a NewDinoflagellate Cyst from the Late Quaternary of the Guaymas BasinGulf of California Mexico Palynology 38(1)101ndash116

Reid PC 1974 Gonyaulacacean dinoflagellate cysts from the British IslesNova Hedwigia 25579ndash637

Rochon A de Vernal A Turon J-L Matthiessen J Head MJ 1999Distribution of Recent dinoflagellate cysts in surface sediments fromthe North Atlantic Ocean and adjacent areas in relation to sea-surfaceparameters American Association of Stratigraphic PalynologistsContributions Series 35146 p

Rochon A Mudie PJ Aksu AE Gillespie H 2003 Pterocysta gen nov Anew dinoflagellate cyst from Pleistocene glacial-stage sediments ofthe black and Marmara Seas Palynology 26(1)95ndash105

Rochon A Lewis J Ellegaard M Harding IC 2009 The Gonyaulax spini-fera (Dinophyceae) ldquocomplexrdquo Perpetuating the paradox RevPalaeobot Palynol 155(1-2)52ndash60

Rossignol M 1962 Analyse pollinique de sediments marins quaternairesen Israeuroel II - Sediments pleistocenes Pollen et Spores 4121ndash148

Rossignol M 1964 Hystrichospheres du Quaternaire en Mediterranee ori-entale dans les sediments Pleistocenes et les boues marines actuellesRevue de Micropaleontologie 783ndash99

Sarjeant WAS 1970 The genus Spiniferites Mantell 1850 (Dinophyceae)Grana 10(1)74ndash78

Schiller J 1935 Dinoflagellatae (Peridineae) in monographischerBehandlung In Dr L Rabenhorstrsquos Kryptogamen-Flora vonDeutschland euroOsterreich und der Schweiz Bd 10(3) Teil 2(2)161ndash320

Soliman A Riding JB 2017 Late Miocene (Tortonian) gonyaulacaceandinoflagellate cysts from the Vienna Basin Austria Review ofPalaeobotany and Palynology 244325ndash346

Sonneman JA Hill DRA 1997 A taxonomic survey of cyst-producingdinoflagellates from recent sediment of Victorian coastal watersAustralia Botanica Marina 40149ndash177

Stein FR v 1883 Der Organismus der Infusionsthiere nach eigenenForschungen in systematischer Reihenfolge bearbeitet II HeuroalfteEinleitung und Erkleuroarung der Abbildungen Leipzig Germany WilhelmEngelmann

Stover LE Evitt WR 1978 Analyses of pre-Pleistocene organic-walled dinofla-gellates Stanford University Publications Geological Sciences 15300 p

Sun Xuekun Song Zhichen 1992 Quaternary dinoflagellates from aren-aceous dolomite in Hainan Island Acta Micropalaeontoligica Sinica9(1)45ndash52 pl 1-2

Seurouto-Szentai M 1982 Szerves vazu mikroplankton biozonak a keuroozep-dunantul pannoniai retegeuroosszleteben Magyar Allami Feurooldtani IntezetEvi Jel Institutum Geologicum Publicum Hungaricum 1980-Rol309ndash344 [in Hungarian]

Shaozhi M 1989 V Dinoflagellata In Hao Yichun Mao Shaozhi RuanPeihua Su Xin Sun Sheping Wang Zhenru Yin Jiayun Zheng Hongeditors Quaternary microbiotas and their geological significance fromnorthern Xisha Trench of South China Sea Wuhan China ChinaUniversity of Geosciences Press p 132ndash147

Shaozhi M Harland R 1993 Quaternary organic-walled dinoflagellatecysts from the South China Sea and their paleoclimatic significancePalynology 17(1)47ndash65

Seurouto-Szentai M 1983 A Pannoniai dinoflagellata egyeurouttesekvizsgalatanak ujabb a-datai euroOslenytani Vitak DiscussionesPalaeontologicae Budapest 2911ndash23 [in Hungarian]

Seurouto-Szentai M 1986 A magyarorszagi Pannoniai (sl) retegeuroosszlet mik-roplankton vizsgalata Folia Comloensis 225ndash45 [in Hungarian]

Seurouto-Szentai M 1988 Microplankton zones of organic skeleton in thePannonian sl stratum complex and in the upper part of theSarmatian strata Acta Botanica Hungarica 34339ndash356

Seurouto-Szentai M 1990 463 Mikroplanktonflora der pontischen (oberpan-nonischen) Bildungen Ungarns Pliozeuroan Pl PontienChronostratigraphie und Neostratotypen p 842ndash869 Jazu and SanuZagreb and Belgrade Yugoslavia [in German]

Seurouto-Szentai M 2000 Organic walled microplankton zonation of thePannonian sl in the surroundings of Kaskantyu Paks and Tengelic(Hungary Annual Report of the Geological Institute of Hungary 1994-1995 2153ndash175) [in Hungarian]

Thurow J Moullade M Brumsack H-J Masure E Taugourdeau-Lantz JDunham K 1988 35 The CenomanianTuronian boundary event(CTBE) at Hole 641A ODP Leg 103 (compared with the CTBE intervalat site 398) In Boillot G et al editors Ocean Drilling ProgramScientific Results Proceedings Leg 103 p 587ndash634

Turon J-L Londeix L 1988 Les assemblages de kystes de Dinoflagellesen Mediterranee occidentale (Mer drsquoAlboran) Mise en evidence delrsquoevolution des paleoenvironments depuis le dernier maximum glaci-aire Dinoflagellate assemblages in the western Mediterranean(Alboran Sea) Evidence of the evolution of palaeoenvironments sincethe last glacial maximum Bulletin des Centres de recherches explor-ation-production Elf-Aquitaine 12314ndash344

Van Nieuwenhove N Potvin E Heikkileuroa M Pospelova V Mertens KMasure E Kucharska M Yang EJ Chomerat N Zajaczkowski M 2018Taxonomic revision of Spiniferites elongatus (the resting stage ofGonyaulax elongata) based on morphological and molecular analysesPalynology 42(Suppl1) httpdoiorg1010800191612220181465736

Wall D 1965 Modern hystrichospheres and dinoflagellate cysts from theWoods Hole region Grana Palynologica 6(2)297ndash314

Wall D 1967 Fossil microplankton in deepndashsea cores from theCaribbean Sea Palaeontology 1095ndash123

Wall D 1971 Biological problems concerning fossilizable dinoflagellatesGeoscience and Man 31ndash15

Wall D Dale B 1967 The resting cysts of modern marine dinoflagellatesand their palaeontological significance Rev Palaeobot Palynol 2(1-4)349ndash354

Wall D Dale B 1968 Modern dinoflagellate cysts and evolution of thePeridiniales Micropaleontology 14(3)265ndash304

Wall D Dale B 1970 Living hystrichosphaerid dinoflagellate spores fromBermuda and Puerto Rico Micropaleontology 16(1)47ndash58

PALYNOLOGY 43

Wall D Dale B 1971 A reconsideration of living and fossil PyrophacusStein 1883 (Dinophyceae) J Phycol 7(3)221ndash235

Wall D Dale B Harada K 1973 Descriptions of new fossil dinoflagellatesfrom the Late Quaternary of the Black Sea Micropaleontology 19(1)18ndash31

Wall D Dale B Lohmann GP Smith WK 1977 The environmental andclimatic distribution of dinoflagellate cysts in modern marine sedi-ments from regions in the North and South Atlantic Oceans and adja-cent areas Marine Micropaleontology 2121ndash200

Warny S 1999 Mio-Pliocene palynology of the Gibraltar Arc a new per-spective on the Messinian Salinity Crisis PhD thesis UniversiteCatholique de Louvain Faculte des Sciences 305 p

Warny SA Wrenn JH 1997 New species of dinoflagellate cysts from theBou Regreg core a Miocene-Pliocene boundary section on theAtlantic coast of Morocco Rev Palaeobot Palynol 96(3-4)281ndash304

Wetzel O 1933 Die in organischer Substanz erhaltenen Mikrofossiliendes baltischen Kreide-Feuersteins mit einem sedimentpetrographi-schen und stratigraphischen Anhang Palaeontogr Ser A 77141ndash188

Williams GL Fensome RA Miller MA Sarjeant WAS 2000 A glossary ofthe terminology applied to dinoflagellates acritarchs and prasino-phytes with emphasis on fossils 3rd ed American Association ofStratigraphic Palynologists Contributions Series 37370 p

Wrenn JH Kokinos JP 1986 Preliminary comments on Miocene throughPleistocene dinoflagellate cysts from De Soto Canyon Gulf of MexicoAmerican Association of Stratigraphic Palynologists ContributionsSeries 17169ndash225

Yunyun Z Morzadec-Kerfourn M-T 1994 Nouveaux kystes dedinoflagelles Spiniferites pacificus nov sp et Pentadinium netangeinov sp de Pleistocene du nord-ouest Pacifique Geobios 27(3)261ndash269

Zhichen S Xueting G Zengrui L Yahui Z Weiming W Zhongheng H1985 A research on Cenozoic palynology of the Longjing structuralarea in the Shelf Basin of the East China Sea (Donghai) regionCenozoic-Mesozoic Palaeontology and Stratigraphy of East ChinaSeries 1 p1ndash209 Anhui Science and Technology Publishing HouseChina [In Chinese with English summary]

44 K N MERTENS ET AL

• Abstract
• • Introduction
  • Discussion of taxa belonging to the genera Achomosphaera and Spiniferites occurring in Pliocene to modern sediments
  • • Achomosphaera andalousiensis Jan du Chne 1977 emend Jan du Chne and Londeix 1988
    • Achomosphaera argesensis Demetresu 1989 (Plate 1 figures 16)
    • Achomosphaera callosa Matsuoka 1983 (Plate 2 figures 13)
    • Achomosphaera granulata Mao Shaozhi 1989
    • Achomosphaera ramosasimilis (Yun Hyesu 1981) Londeix etal 1999
    • Spiniferites alaskensis Marret etal 2001 ex Marret in Fensome Williams 2004
    • Spiniferites asperulus Matsuoka 1983 (Plate 3 figures 16)
    • Spiniferites belerius Reid 1974
    • Spiniferites bentorii (Rossignol 1964) Wall Dale 1970
    • Spiniferites bulloideus sensu Wall (1965)
    • Spiniferites coniconcavus De Schepper etal 2004
    • Spiniferites cruciformis Wall and Dale in Wall etal 1973
    • Spiniferites delicatus Reid 1974
    • Spiniferites elongatus Reid 1974
    • Spiniferites falcipedius Warny Wrenn 1997
    • Spiniferites firmus Matsuoka 1983 (Plate 3 figures 711)
    • Spiniferites hainanensis Sun Xuekun Song Zhichen 1992
    • Spiniferites hexatypicus Matsuoka 1983 (Plate 4 figures 14)
    • Spiniferites hyperacanthus (Deflandre Cookson 1955) Cookson Eisenack 1974
    • Spiniferites lazus Reid 1974
    • Spiniferites ludhamensis head 1996 (Plate 4 figures 516)
    • Spiniferites membranaceus (Rossignol 1964) Sarjeant 1970
    • Spiniferites mirabilis (Rossignol 1964) Sarjeant 1970
    • Spiniferites multisphaerus Price Pospelova 2014
    • Spiniferites nanus Matsuoka 1976
    • Spiniferites nodosus (Wall 1967) Sarjeant 1970 (Plate 5 figures 13)
    • Spiniferites pachydermus (Rossignol 1964) Reid 1974
    • Spiniferites pacificus Zhao Yunyun Morzadec-Kerfourn 1994
    • Spiniferites pseudofurcatus subsp obliquus (Wall 1967) Lentin Williams 1973 (Plate 5 figures 47)
    • Spiniferites ramosus sensu Rochon etal (1999)
    • Spiniferites ramosus subsp multiplicatus (Rossignol 1964) Lentin Williams 1973
    • Spiniferites rhizophorus Head in Head Westphal 1999 (Plate 6 figures 115)
    • Spiniferites ristingensis Head 2007 (Plate 7 figures 115)
    • Spiniferites scabratus (Wall 1967) Sarjeant 1970 (Plate 8 figures 16)
    • Spiniferites septentrionalis Harland 1977 (Plate 8 figures 79)
    • Spiniferites spinatus (Song Zhichen in Song Zhichen etal 1985) Lentin and Williams 1989
    • Spiniferites strictus Matsuoka 1983 (Plate 8 figures 1012)
    • Spiniferites tripodes (Morzadec-Kerfourn 1966) Lentin Williams 1973
    • • Species previously assigned to Spiniferites but here considered or accepted as belonging to other genera
      • • Impagidinium inaequalis (Wall Dale in Wall etal 1973) Londeix etal 2009 (Plate 9 figures 16)
        • Spiniferites rubinus (Rossignol 1962 ex Rossignol 1964) Sarjeant 1970
        • Thalassiphora balcanica Balte 1971
        • • Discussion about genusspeciesvariety concepts
          • Acknowledgements
          • References