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Nesticus eremita (Araneae: Nesticidae): redescription of a potentially invasive European spider found in New Zealand PDF

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2011.TheJournalofArachnology39:511-514 SHORT COMMUNICATION Nesliciis eremita (Araneae: Nesticidae): redescription ofa potentially invasive European spider found in New Zealand CorJ. Vink'-^and NadineDuperre-"*: 'Biosecurity Group, AgResearch, Private Bag4749, Christchurch 8140, New Zealand. E-mail:[email protected];-EntomologyResearchMuseum,POBox84, LincolnUniversity, Lincoln 7647, NewZealand; -''Division ofInvertebrateZoology, American Museum ofNatural History, Central Park West at 79th Street, NewYork, NewYork 10024 USA Abstract. NesticuseremitaSimon1879isnaturallyfoundincavesinsouthernEurope.Ithasalsoinvadedandestablished itselfin Germanyand has now been found in an abandoned air-raid tunnel in Auckland, New Zealand. A diagnosis, redescription,fullsynonymyandillustrationsarepresentedtoaidintheidentificationofthispotentiallyinvasivespider. Keywords: Cavespider,invasivespecies,taxonomy,troglophile AnumberofspeciesinthefamilyNesticidaehaveinvadedareas ItseemsalmostcertainthatN. eremitaarrivedinAucklandviaa outside their natural range. Nesticus cellulanus (Clerck 1757) is shippingcontainer, asthe tunnel entrance inAlten Reserve isonly originallyfromEurope,butisnowfoundinthenortheasternUSA 600metersfromthePortofAuckland,whichisoneofNewZealand’s ‘ a20n0d9)Quebec (Gertsch 1984; Paquin & Duperre 2003; Paquin & busiest seaports. The air-raid tunnels beneath Albert Park in Hedin 2005). Eidmamtellapallida (Emerton 1875) was originally Auckland city are extensive and were partially filled with unfired from North and Central America (Gertsch 1984), but is now bricks at the end of the Second World War, when most of the found throughout the world (Platnick 2011). Nesticella mogera entranceswereblockedoff(Pilkington2008). (Yaginuma1972),isnativetoAsiabutintroducedtoFiji,Germany The male specimen collected had built a web and was in good and Hawaii (Lehtinen & Saaristo 1980; Gertsch 1984; Kielhorn condition. It would be extremely unlikely that the single male . collectedhadlived longenoughtostowawaywithcargo,madethe Here we document another nesticid that has spread outside its journeyfromEuropetoNewZealand(ittakesabout40daysfora naturalrange.NesticuseremitaSimon 1879occursnaturallyincaves cargo ship to sail from Italy to Auckland (M.R. McNeill pers. 2t0h1r0o)ughout Italy, southeast France, Corsica, Switzerland, Slovenia, comm.)),walkedthe600mfromtheporttothetunnelandthenmade Croatia,Montenegro,Bosnia-HerzegovinaandGreece(Lessert1906, a web. It is more likely that the male was part ofan established 1910;Kratochvil 1933;Dresco 1966;Dresco&Hubert 1967;Brignoli populationor,attheveryleast,theoffspringofagravidfemalethat 1971, 1977;Deeleman-Reinhold 1974;Maurer&Hanggi 1990). Itis did live long enough to make the journey from Europe. The I also found in cave-like synanthropic habitats (e.g., tunnels, cellars, establishmentofN. eremitainAucklandneedsfurtherconfirmation. catacombs,railroadtrackballast,wells)initsnaturalrange(Brignoli Unfortunately, the entrance to the tunnel where the single male 1971; Maurer& Hanggi 1990)and where it isadventivein Austria specimenofN. eremitawascollectedin2000hassincebeenblocked (Knoflach &Thaler 1998) and Germany (Jiiger 1995, 1998; Staudt off,soitwasnotpossibletosearchforfurtherspecimensandconfirm .Nesticuseremitahasestablishedpopulationsinsewertunnelsin whetherthereisstillapopulationthere. the German cities ofCologne, Mainz and Mannheim (Jager 1995, Given that people generally do not spend a great deal oftime [ 1998).ThesecitesareallontheRhineRiver,andN.eremitamayhave lookingforsmallspidersinsewersandabandonedtunnels,itisquite I spread downstream from its endemic range in the Alps either possiblethatN. eremitahasspreadtootherpartsoftheworldand naturallyorviahumantransport(Jager 1998).Giventhatthesewer established undetected populations. The presence ofN. eremita in I tunnels in which N. eremita was found were not much more than human-madetunnelsin Auckland, Cologne, Mainzand Mannheim 100yearsold(Jager1995, 1998)andthatthespiderfaunaofCentral doesnotpresentanimmediatethreattonaturalecosystems.However, ' Europeiswellknown(e.g.,Heimer&Nentwig 1991),itappearsthat ifthisspeciesweretospreadtonaturalcavesystemsoutsideitsnative N. eremita has become established in Germany comparatively range, it has the potential for making ecological impacts. Invasive recently. spider species can harm endemic spiders through competitive On21 June2000,CJVandGraceHallwenttoanabandonedair- displacement (Nyffeler et al. 1986; Hann 1990; Bednarski et al. raid tunnel in Alten Reserve (36°51.04'S, 174°46.42'E) in central 2010),and invasivearthropod predatorscan impact nativecommu- AucklandinsearchoftheAustralianstiphidiidProcamhridgeagrayi nities(Snyder&Evans2006). Thismaybeeven moreofaconcern I Davies2001,whichhadbeencollectedfromtherein 1999(Davies& in cave ecosystems, which may be more at risk from ecological Lambkin2001). NospecimensofP. grayiwerefound, butasingle disturbance(Reeves2001). maleN. eremitawascollected from aweb neartheentranceofthe The species descriptions of N. eremita are all in relatively II tNuensnteilc.idaNee,itahserthactolflaemctiolryhreacdognnoitzebdeetnherecsopredceidmefnroamsNbeelwonZgeianlgantdo diniaacgcneossstiibcleillpuusbtlriactaiotnisonosf,mnaolte ainndEfnegmlailsehgeonritaldioa tnoogtethefre.atuTroe I (Paquinetal.2010;Sirvid,etal.2011).Thespecimenwaspreservedin facilitate the identification ofthis potentially invasive species we 70% EtOH, labeled and put amongst the unsorted material at the present a full synonymy, diagnosis, brief redescription and NewZealandArthropodCollection(NZAC);itremainedthereuntil illustrations ofthe habitus, male pedipalp and female epigynum. 1 November2010whenCJVnoticeditwhilesearchingforspecimensof Terminologyofthemalepedipalpal structures followsAgnarsson Mimetidae. etal. (2007). j 511 — — 512 THEJOURNALOFARACHNOLOGY Figures !^. Nesticuseremita,femalefromMannheim,malefromMainz,Germany. 1.Habitus,female;2.Maleleftpedipalp,ventralview, tta = theridioidtegularapophysis,e = embolus, ma = median apophysis, t = tegulum, st = subtegulum,pc = paracymbium; 3. External genitalia,ventralview,co =copulatoryopening,ms = mediumseptum;4.Internalgenitalia,dorsalview. TAXONOMY NesticuseremitaitalicaCaporiacco 1934:401,figs.2,3(Synonymized byDresco&Hubert 1967). NesticidaeSimon 1894 NesticusspeluncarumeremitaKratochvil 1933:40,63,figs. 3, 13, 14, | NesticinaeSimon 1894 27-30; Wiehle 1963:435, figs. 8-10; Wiehle 1967:193, figs. 45,46 NestNiecsutsiceursemTihtoareSlilmo18n691879 Ive(siSaynoerneymmiitzaedLebhytiDnreensco&&SHauabriesrtto1916978)0.:51 (Transferred from (Figs. IM) Nesticus). NesticuseremitaSimon 1879:258;Simon 188:48,pi.26,fig.8;Lessert Materialexamined.—NEWZEALAND:Auckland,AltenReserve, 1906:610,pi.20,figs. 3-5; Lessert 1910:300,figs. 164-166;Simon 21 June 2000, C.J. Vink &G. Hall leg, Id(NZAC). GERMANY: 1929:658,753,figs. 1015,1016;Dresco1966:805;Dresco&Hubert Mannheim,24November1997,P.Jagerleg. Id19(AMNH);Mainz, 1967:3; Brignoli 1971:206, figs. 118-122; Deeleman-Reinhold 2! May 1996,P.Jagerleg. Id 19(AMNH). 1974:12, fig. 9; Brignoli 1975:28; Kratochvil 1978:39, figs. 3, 4; Diagnosis. Nesticus eremita can be distinguished from other Thaler 1981:274,figs. 6,8;Heimer&Nentwig 1991:276,fig. 732; Nesticidaebyfeaturesofthemalepedipalp(Fig.2),particularlythe Jager 1998:13,fig. lA. structureofthetheridioidtegularapophysis,theshapeofthelarge, Theridion parenzani Trossarelli 1931:13, fig. 1 (Synonymized by unbranchedparacymbiumandthestructureoftheexternalgenitalia Brignoli 1975); of the female (Fig.3). The theridioid tegular apophysis is more NesticusstrasseriRoewer 1931:14, fig. 11, 13c, d (Synonymized by compact than that of N. cellulanus, and the distal end of the Kratochvil 1933). paracymbium is rounded. The slit-like copulatory openings ofthe — VINK&DUPERRE—POTENTIALLYINVASIVENESTICUSEREMITA 513 externalgenitaliadivergeanteriorlyandthemediumseptumnarrows informationontimestaken bycargoshipsto reachAuckland. We posteriorly, whereas in N. celliilaniix, the copulatory openings thank Norman Platnick (American Museum ofNatural History), conRvedeersgceriapnttieorni.or—lyC,olaonrdtihnemalecdoihoulm:sMeaplteumcisarwaipdaeceposltiegrhitorlyye.llow; IrnegviiewAegrnsarfsosronco(mUmnievnetrssitoynofthPeuemratnousRcirciop)t.anCdJVtwowaasnopnarytmioalulsy abdomen off-white; legs yellow, metatarsi, tarsi oflegs I, II light funded by New Zealand’s Foundation for Research, Science and orange.Femalecarapacelightyellowwithblacktrident-shapedmark Technology through contracts C02X0501 (Better Border Biosecu- extendingfromfoveatolateraleyes(Fig. 1);abdomenoff-whitewith rity (B3) programme (www.b3nz.org)) and C09X0501 (Defining darkpattern(Fig. 1);legsyellow-orangewithdarkbands(Fig. 1). New Zealand’s Land Biota / Te Tautuhi i nga Hanga Koiora o Malepedipalp(Fig.2):theridioidtegularapophysiscompact,with Aotearoa). several pointed projections; paracymbium long, unbranched with LITERATURECITED rounded end. Male chelicerae with 3 promarginal teeth, 5 retro- marginaldenticles. Agnarsson, I., J.A. Coddington & B. Knoflach. 2007. Morphology Epigynum (Fig.3)withslit-likecopulatoryopeningsthatdiverge andevolutionofcobwebspidermalegenitalia(Araneae,Theridi- anteriorly, medium septum narrow posteriorly; internal genitalia idae).JournalofArachnology35:334-395. (Fig.4) with branched spermathecae. Female chelicerae with 3 Bednarski,J.,H.Ginsberg&E.M.Jakob.2010.Competitiveinterac- promarginalteeth, 10retromargina!denticles. tions between a native spider {Frontinella communis, Araneae: Dimensions (mm). Male (Auckland): total length 3.5; carapace Linyphiidae)andaninvasivespider[Linyphiatriangularis,Araneae: length 1.9, width 1.8, height 0.8; abdomen length 2.6, width 1.4; Linyphiidae).BiologicalInvasions12:905-912. sternumlength1.1,width1.1;totallengthoflegI16.7,legII11.7,leg Bosselaers,J. 1998.Nesticushenderickxi(Araneae,Nesticidae),anew III8.7,legIV11.9.Female(Mainz):totallength5.2;carapacelength blindtroglobiticspiderfromCrete. BulletinoftheBritishArach- 2.1, width 1.7, height 0.9; abdomen length 3.3, width 2.3; sternum nologicalSociety 11:9-14. length,1.1,width1.1;totallengthoflegI 15.8,legII 10.9,legIII7.7, Brignoli, P.M. 1971. Note su ragni cavernicoli italiani (Araneae). legIV 11.2.— FragmentaEntomologica7:121-229. Variation. Female:Carapacelength 1.7-2.6;totallength3.8-5.4. Brignoli,P.M. 1975.Ragnid’ltalia.XXV.Sualcuniragnicavernicoli Male: Carapace length 1.7-2.3; total length 3.5-4.7. Based on deiritaliasettentrionale(Araneae).NotiziariodelCircoloSpeleo- materialexaminedandJager(1998). Thedegreeofpigmentation in logicoRomano20:7-39. N.eremitacanvary(Jager1998);weobserveduniformlylightcolored Brignoli, P.M. 1977. Ragni de Grecia X. Nuovi dati sulla Grecia specimens and specimens with a contrasting color pattern on the continentale ed insulare (Araneae). Revue Suisse de Zoologie abdomenandlegs(Fig. 1). 84:937-954. Distribution. Nesticuseremitaisfoundincavesinitsnaturalrange Caporiacco, L. 1934. INesticusliguriedemiliani.AnnalidelMuseo throughout Italy, southeast France, Corsica, Switzerland, Slovenia, CivicodiStoriaNaturalediGenova56:395-405. Croatia, Montenegro, Bosnia-Herzegovina and Greece. It has also Davies, V.T. & C.L. Lambkin. 2001. A revision ofProcamhridgea beenfoundinsynanthropichabitats(e.g.,tunnels,cellars,catacombs, Fortser&Wilton,(Araneae:Amaurobiodea: Stiphidiidae). Mem- railroadtrackballast,wells)inItaly,Switzerland,Austria,Germany oirsoftheQueenslandMuseum46:443^59. andNewZ—ealand(Auckland). Deeleman-Reinhold,C.L.1974.ThecavespiderfaunaofMontenegro Biology. Like most nesticids, Nesticus eremita is a troglophile. (Araneae). Glasnik Republickog Zavoda za Zastitu Prirode i SpecimensofN. eremitathatlivewellawayfromsunlighthaveless PrirodnjackogMuzejaTitogradu6:9-33. somDaNtiAcpsieqgumeenncteast.i—onDaNnAdrseedquuceendcetsapfertaom(Jtahgeermi19t9o8c)h.ondrial genes DreNsecsot,icEu.s(1f9a6m6..NEetstuidceidadee).quAenlnqauleessedsepeScpeesiedo’laorgaiieg2n1ee:s795d-u81g3e.nre cEyUt7o4c6h4r3o6m)eacnodxi1d6aSserisbuobusnoimtal1R(NCOAI)((EGUe74n6B4a4n5k)awcecersesiroenponrutemdbeirn DreNsecsot,icEu.se&reMmi.taHSubiemrotn.(A19r6a7n.eaEet,uNdeestidceisdavea)r.iaAtricohnisveoscudleaiZroeoslocghieez Lopez-Panc—orbo&Ribera(2011). ExperimentaleetGenerale108:3-31. Remarks. Thepositionofthetrichobothriumonmetatarsus 1 has Gertsch,W.J. 1984.ThespiderfamilyNesticidae(Araneae)inNorth beenpromotedasanimportantcharacterfordistinguishingbetweenN. America, Central America, and the West Indies. Bulletin ofthe eremita&ndN.cellidcmus(WiehXe.1963;Thaler1981;Heimer&Nentwig TexasMemorialMuseum31:1-91. 1991)and forhighernesticid taxonomy(Lehtinen & Saaristo 1980); Hann,S.W. 1990.EvidenceforthedisplacementofanendemicNew however, Jager (1998) found that this character was not diagnostic Zealand spider, Latrodectuskatipo Powell by the South African between N. eremita and N. celhikmus. Lehtinen & Saaristo (1980) species Steatoda capensis Hann (Araneae: Theridiidae). New transferredNesticuseremitatothegenusIvesiaPetrunkevitch1925,but ZealandJournalofZoology 17:295-308. Kaston (1945) and Gertsch (1984) considered Nesticus a senior Heimer, S. & W. Nentwig. 1991. Spinnen Mitteleuropas: Ein synonymofIvesia. Thepedipalpal scleritesofN. eremitaaresimilar Bestimmungsbuch. PaulParey, Berlin. tothoseofN.cellulanus,thetypespeciesofNesticus.andappeartobe Jager, P. 1995. Erstnachweis von Holocnemusplucliei und zweiter homologus.Thesetwospeciesalsoappeartobecloselyrelatedbasedon NachweisvonNesticuseremitafiirDeutschlandinKoln(Araneae: a phylogenetic analysis ofmitochondrial DNA (Lopez-Pancorbo & Pholcidae,Nesticidae).ArachnologischeMitteilungen 10:23-24. Ribera 2011). There are European Nesticus species with greater Jager, P. 1998. Weitere Funde von Nesticus eremita (Araneae: morphological similaritiestoN. eremita(e.g., N. spehmcarum Pavesi Nesticidae) in Siiddeutschland mit Angaben zur Taxonomie im 1873,N.henderickxiBosselaers 1998),but unlikeN. cellulanus,these VergleichzuN.cellulanus.ArachnologischeMitteilungen15:13-20. species are only known from their natural cave habitats and have Kaston, B.J. 1945. New Micryphantidaeand Dictynidaewith notes limiteddistributions(Dresco1966;Brignoli 1971;Bosselaers1998). onotherspiders.AmericanMuseumNovitates 1292:1-14. ACKNOWLEDGMENTS Kielhorn,K.-H.2009.FirstrecordsofSpermophorakerinci,Nesticella mogera and Pseudanapis alalia on the European mainland Grace Hall is always an excellent host when visiting the New (Araneae: Pholcidae, Nesticidae, Anapidae). Arachnologische ZealandArthropodCollection. PeterJager(SenckenbergMuseum) Mitteilungen37:31-34. kindlygiftedspecimensofN.eremitatoAMNHforustostudyand Knoflach,B.&K.Thaler.1998.KugelspinnenundverwandteFamilien he and Theo Blick (Senckenberg Museum) provided information von Osterreich: Okofaunistische Ubersicht (Araneae: Theridiidae, about German localities. Thanks to Mark McNeill for providing Anapidae,Mysmenidae,Nesticidae).Stapfia55:667-712. 514 THEJOURNALOFARACHNOLOGY Kratochvil, J. 1933. Evropske druhy celedi Nesticidae Dahl. Prace Management Symposium. (G.T. Rea, ed.). Southeastern Cave MoravskePrirodovedeckeSpolecnosti8(10):1-69. Conservancy,Chattanooga,Tennessee. Kratochvil, J. 1978. Araignees cavernicoles des iles Dalmates. Roewer,C.F. 1931.ArachnoideenaussiidostalpinenHohlengesam- PfirodovednePraceUstavuCeskoslovenskeAkademievedvBrne melt von Herrn Karl Strasser in den Jahren 1929 und 1930. 12(4):l-59. MitteilungenfiberHohlenund Karstforschung 1931:1-17. Lehtinen, P.T. & M.l. Saaristo. 1980. Spiders of the Oriental- Simon, E. 1879. Arachnides nouveau de France, d’Espagne et Australian region. IL Nesticidae. Aiinales Zoologica Feniiici d'Algerie. Premiermemoire. BulletindelaSocieteZoologiquede 17:47-66. France4:251-263. Lessert, R. 1906. Araneae. InCarl,J., BeitragzurHohlenfaunader Simon, E. 1881. Les Arachnides de France. Tome 5. lere partie insubrischenRegion. RevueSuissedeZoologie 14:601-615. contenantlesfamiliesdesEpeiridae(supplement)etdesTheridio- Lessert, R. 1910. Catalogue des invertebres de la Suisse. Ease. 3, nidae(commencement). LibrairieencyclopediquedeRoret,Paris. Araignees. Museed’histoirenaturelledeGeneve,Geneva. Simon, E. 1929. Les Arachnides de France. Synopsis general et Lopez-Pancorbo,A.&C.Ribera.2011.Nesticushaeticussp.n.,anew cataloguedesespecesfrangaisesdeI’ordredesAraneae.Tome6,3e troglobitic spider species from south-west Europe (Araneae, partie,Paris. Nesticidae).ZooKeys89:1-13. Sirvid, P.J., Z.-Q. Zhang, M.S. Harvey, B.E. Rhode, D.R. Cook, Maurer,R.&A.Hiinggi. 1990.KatalogderschweizerischenSpinnen. 1. Bartsch & D.A. Staples. 2011. Chelicerata: horseshoe crabs, DocumentaFaunisticaHelvetiae,Neuchatel,Switzerland. arachnids, sea spiders. Pp. 50-89. In New Zeala—nd Inventory of Nyffeler, M., C.D. Dondale & J.H. Redner. 1986. Evidence for Biodiversity. Volume Two: Kingdom Animalia Chaetognatha, d((HAiersnaptnlzea)ac,ee:mebTnyhteritodhfieidaaEeu)Nr.oorCptaehnaandAimsaepnreicJicoeausnrnaYs.lpiodhfeirZp,iomoSclttoaegtayato6d4(a:L8ib6no7nr-ae8ea7ul4si.)s SnysEdieetdyry,PsroWezs.osE,a.,ChI&rcihsnEto.fcWohs.usriclEhsv,.anN(sDe..wP.2Z00eG6ao.lradnEodcn.o,loegdi.c)a.lCaefnfteectrsbuorfyiUnnviavseirv-e Paquin,P.&N.Duperre.2003.Guided’identificationdesAraignees arthropod generalist predators. Annual Review of Ecology, (Araneae)duQuebec.Fabreries.Supplement 11:1-251. Evolution,andSystematics37:95-122. Paquin,P.&M.Hedin.2005.Nesticidae.Pp. 178-180.InSpidersof Staudt, A. 2010. Nachweiskarten der Spinnentiere Deutschlands NorthAmerica:AnIdentificationManual.(D. Ubick,P. Paquin, (Arachnida: Araneae, Opiliones, Pseudoscorpiones). Deutsche P.E. Cushing & V.D. Roth, eds.). American Arachnological Arachnologische Gesellschaft, onlineat http://www.spiderling.de/ Society,Keene,NewHampshire. arages. PaqALuininnnco,otlaPn.t,,eNCd.eJF.wamVZiielnayklaK&ned.yN.anDdupSeprercei.es20L1is0t..SMpaindearaskiofWNheewnuZaeaPlraensds:, TTrhoasAlsreaarr,neellaKie.,:1FN9.e8s1t1.i9c3iF1di.abeeA)rr.aNcSenesintdciikcuesnrabciecdrorglitaiciaininsaelRBliaooelGworegoirtctaa19631d1i:27P(1Ar-ro2am7co6hn.ntiodrae: PPilalsMtkhniueinlscgtktee,orusnN.,m.ALoSr.f2c0hN21aa01te0.uo8rTl.aohlgeHyeHWriioisnrttaolNrgdeye,wSpvNiaZedleeuwarelsaCYanootdrfakl5.ot1gh:O,e1n0lv6Aie-lrn1bse1ei7roa.ntth1Pt1ta.p5r.:k/A/mraeeisrreiarrccaahin.d WWiieeA2(hhI1rlls:atee1nr,,i3ea-a)H1e.H4.)...At1tS9i1e69n76d.c3ek.leMlne’/bUAfebclre,cg-raiediaennmNeaeisasBteiiSmocciluieosengnittceihafloierc4gua4yt:nzV4eke3yn1iGe-at4to3R-t5eT.uirnmegonstdieenrro-IA(sArtraraniceahan.ea, amnh.org/entomology/spiders/catalog/INTRO1.html (Arach.). SenckenbergianaBiologica48:183-196. Reeves, W.K. 2001. Exotic species in North American caves. Pp. 164-166.InProceedingsofthe 1999NationalCaveandKarst Manuscriptreceived2July2011, revised25September2011.

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