Description of the female of Orsolobus pucara Forster & Platnick 1985, with comments on the functional morphology of the female genitalia in Dysderoidea (Araneae: Dysderoidea: Orsolobidae)

2010·7.7 MB·English

by  Matías Andrés Izquierdo

2010.TheJournalofArachnology38:511-520
DescriptionofthefemaleofOrsolohuspucaraForster& Platnick 1985,withcommentsonthefunctional
morphology ofthe female genitalia in Dysderoidea (Araneae: Dysderoidea: Orsolobidae)
MatiasAndresIzquierdoand FacundoMartinLabarque: Division ofArachnology, MuseoArgentino deCiencias
Naturales “Bernardino Rivadavia”, AvenidaAngel Gallardo470 C1405DJR, BuenosAires, Argentina. E-mail:
[email protected]
Abstract. The female ofOrsolohuspucara Forster& Platnick 1985 is described and its genitalia examined using the
scanningelectronmicroscope(SEM).Asmallphylogeneticmatrixwithfemalegenitalandsexualbehaviorcharacterswas
madewiththeaimtostudytheevolutionofthesecharactersinthesuperfamilyDysderoidea.Thisisthefirsttimethatthe
femalegenitaliaofaspeciesofthefamilyOrsolobidaehavebeenstudiedindetailwithSEM.Theanteriorportionofthe
femalegenitaliaisasclerotizedstructurewithglandductsandsitesformuscleattachments.Theposteriorportionhasa
membranousreceptaculumandasclerotizedplatethatservesasattachmentformuscles.Wediscusstheprobablefunction
of genital characters in a phylogenetic context. The anterior sclerotized elements of the female genitalia of some
Dysderidae,OrsolobidaeandOonopidaespeciesandtheanteriorreceptaculumintheSegestriidaeseemtobehomologous
structuresbecauseofthepresenceofglandductsandsperm.However,bothofthesecharacteristicsarelostinsomespecies
ofthesefamilies,theanteriorportionofthefemalegenitaliabeingtransformedintoahighlymodifiedstructureserving
mainlyasattachmentformusclesimplicatedinsexualbehaviormechanisms.
Keywords: Characterevolution,complexgenitalia,reproductivebehavior,spiders,taxonomy
The family Orsolobidae Cooke is a group of haplogyne fosuma Burger 2002 (probably to be transferred to another
spiderswithsixeyesthatcanbydistinguishedbythepresence genusinthefuture). Fordetailscomparefig. 2binGrismado
ofan elevated tarsal organ (Fig. 7D). These active hunting 2008withfig. 3 in Burgeretal. 2003).
spidersaredistributedineasternandwesternAustralia, New WiththisworkwewishtoprovidethefirstSEMimagesof
Zealand,SouthAfrica,Argentina,FalklandIslands,Chileand the female genitalia in the family. Also, we compare the
Brazil (Forster & Platnick 1985; Griswold & Platnick 1987; morphology ofthe female genitalia of O. pucara with other
Platnick&Brescovit 1994; Brescovitelai. 2004; Baehr2009). species of Dysderoidea. We used published data about the
TheOrsolobidae,togetherwiththeDysderidae,Oonopidae functionalmechanismsofthegenitaliaacrossthesuperfamily
and Segestriidae, are grouped in the haplogyne superfamily to infersimilarpatterns in O. pucaraand theOrsolobidae in
Dysderoidea by the occurrence of a second portion of the general, to detect possible homologies and to discuss the
internalfemalegenitaliaassociatedwiththeposteriorwallof evolutionofthefemalegenitalcharacters. Detailedimagesof
the bursal cavity (Forster & Platnick 1985; Ramirez 2000). the male palp are also presented, and other anatomical
Althoughthefemalegenitaliaofmanyhaplogynespidersare structuresofthefemaleareillustrated.
simple,suchasintheFilistatidaeandCaponiidae,thegenital
structuresofotherfamilies(amongthemtheOrsolobidaeand METHODS
Oonopidae)appearrathercomplex(Burger&Kropf2007).In Specimensaredepositedinthecollectionofarachnidsofthe
somespeciestheanteriorsectionofthefemalegenitalia(AFG Museo Argentino de Ciencias Naturales “Bernardino Riva-
henceforth) has a very complex organization. It has been davia”, BuenosAires(MACN-Ar, CristinaScioscia).
proposed that the degree of complexity might involve The format ofdescriptions and morphologic terminology
mechanisms of cryptic female choice, sperm dumping, and followsingeneralForster&Platnick(1985). Indescribingthe
genital organization similarto the entelegynecondition (Uhl femalegenitalia,weusedthecriterionfollowedbyPlatnicket
2000;Huber2002;Burgeretal.2003,2006;Huberetal.2005; al. (1999) for naming structures situated anteriorly or
Burger 2007; Burger & Kropf 2007). The oonopid genus posteriorly to the uterus externus. Abbreviations used for
ScaphiellaSimon 1891 isinfactfunctionallyentelegyne,since eyesandlegsarestandardinarachnology. Measurementsare
theyhave separatecopulatoryand fertilization openings and in millimeters. After dissection, the female genitalia were
ducts (see Burger 2009). The same condition occurs in the digestedinhotKOHandmountedintemporarypreparations
diversegenusEscaphiellaPlatnick&Duperre2009(Platnick& with lacticacid. The malepalpwascleared withcloveoil. A
Duperre2009). cameralucidamountedonacompoundmicroscope(Olympus
AlthoughForster&Platnick(1985)illustratedthediversity BH-2)wasusedtomakedrawings. Photographsofpreserved
of female genital structures in the Orsolobidae, the fine spiders were made with a digital camera (Nikon DXM1200)
structure of this group is unknown, thus precluding more mounted on a stereoscopic microscope (Nikon SMZ 1500).
detailed functionalhypotheses. Also, homologiesaredifficult The focal planes were combined with Helicon Focus 3.10.3
toexplorewhencomparing,forexample,thesimplegenitalia (onlineathttp;//helicon.com.ua/heliconfocus/). Scanningelec-
ofsegestriid genera like Segestrki Latreille 1804 or Ariadna tronmicrographsweretaken underhighvacuum witha FEI
Audouin 1826 with the complex configurations found in XL30TMPaftercritical pointdryingandAu-Pdcoating. A
oonopids like Antoonops Fannes &Jocque 2008 or Opopaea small phylogenetic matrix with genital and sexual behavioral
511
512 THEJOURNALOFARACHNOLOGY
—
Table 1. Terminalincludedinthephylogeneticanalysisandthedatasourcewherethecharacterswereconstructed.
Family Terminal Datasource
Caponiidae NopsMacLeay 1839 Izquierdo&Labarquepers.obs.
Segestriidae Ariadnahoesenbergi Grismado2008;Izquierdo&Labarquepers.obs.
Dysderidae Dysderaerythrina Uhl,2000
Hapactealepida(C.L.Koch 1838) Burger&Kropf2007
Orsolobidae Orsolobuspucara Forster&Platnick 1985;Izquierdo&Labarquepers.obs.
OsornolobusForster&Platnick 1985 Forster&Platnick 1985
Oonopidae ScaphiellaliesperaChamberlin 1924 Burger2009
AntoonopscorbiiloFannes&Jocque2008 Fannes&Jocque2008
Silhouettellaloricatula(Roewer1942) Burgeretal.2006
Opopaeafosuma Burgeretal.2003
Orchestina(sp.l) Izquierdo&Labarquepers.obs.;Burgeretal.2010
Orchestina(sp.2) Izquierdo&Labarquepers.obs.
Grymeus Burger2010
Lionneta Burger2010
Myrmopopaea Burger2010
characters includes morphological and behavioral characters LegI(Fig. 7D)andabouttencuticularlobesandone(maybe
describedintheliteratureandfromourpersonalobservations two) receptoron leg IV. Retroclaw and proclawwith fifteen
(Fig. 8). Theterminalsand sourcesarelistedinTable 1. The teeth on both outer and inner margins (Fig. 1C). Tricho- '
phylogenetic tree was taken from the previous analyses of bothrial socket with proximal hood at the same level as the
Platnicketal.(1991)andRamirez(2000).TheOonopidaewas cuticleandwiththesamesculpture(Fig. 7E).Distalhoodvery
considered monophyletic, butwithoutany internal structure, short and with same sculpture as cuticle. Base of the
i
except for two groups supported by potential evidence: the trichobothrial seta slightly swollen and with oblique rings
Lionneta Benoit 1979, Grymeiis Harvey 1987 and Myrmopo- (Fig. 7E). AFG heavily sclerotized, formed by only one [
paea Reimoser 1933 clade (see Burger 2010) and the genus anterior median plate (mp. Fig. 3A) with four basal spurs,
mOracphpeestdinoanStihmisontre1e88u2si(njgumTpNinTg(oGoonolpoibdosf)f.eCthaalr.ac20t0e8r)s.wTehree atrwioseosftthheemandtierreicotredmedodrisaanllyroadnd(mtrw)o,vwehntircahllyb.eaBrestwneuemnertohuesm ^
aim ofthis small analysis is to explore the evolution ofthe gland ducts near its base (Figs. 3C, 4C, E). The tip ofthe 'j
female genitalia characters in the Dysderoidea; a full medianrodisbifurcatedandhasseveralscarscorresponding
rtheiasnacloynstirsibouftidoyns.deroid relationships is beyond the scope of rtoodthheasplaacfelasttoefnemduspcrlojeesctaitotnacdhirmeecnttesd(vFeingt.ra4lDl)y.thTahtemameydailasno ij,
bearmuscleinsertions(Fig. 3B). Posteriorpartofthefemale
SYSTEMATICS genitalia (PFG) with a membranous posterior receptaculum ';!
Orsolobus—pucar(aFiFgosr.st1e-r7)&Platnick 1985 FfBioegrt.mwee3deB)nbtythheaamttuambreea-yltiwkoeacstscelceatrsiootniszutephdpaoptlratetinendsgs(Fisintgr.auc3stAau,crkeasssttefrrouicrstkurtoehn.e f
Femalediagnosis. Easilydistinguishedfromotherfemales receptaculum oras attachments formuscles that control the I
ofthegenusbytheshapeofthemedianrod,bifurcatedatthe aperture of these structures. External surface of the sack
tipand with a flattened projectiondirected ventrally(Figs. 3 structure withmanygland ducts formed by short bases(BS) I
;
A,DBe)s.cription.—(MACN-Ar 16120). Total length 3.47, cara- aspnadrsdeilsytaldipsitrriifbourtmedcaoprs(gDrCo)up(eFdig.in5A)t.woTheorgltahnrdeeduoctnsatrhee •^
pace 1.40wide,opisthosoma 1.80wide. LeglengthI: 6.49, II: receptaculum surface and communicate into the lumen
5.90, III: 4.93, IV: 6.44; palp length 2.08. Carapace pale through simple pores (Figs. 5A, B). There is a “posterior
orange with several setae in the surface (Fig. 1); legs and plate” (pp, Figs. 3A, 4A) incloseconnection withtheAFG. |,
maxillary endites pale yellow, sternum and labium pale Theposteriorplatehasaconvexshapeinthemedianlineand
orange. Opisthosoma pale yellowish with many dots of extends toward both sides, acquiring a flattened shape (fa,
pigment, visible by transparency through the cuticle “flattened lateral apodemes”. Fig.4B). The position ofthe Ij,
(Figs. lA-C). Spinnerets yellow. ALE and PLE contiguous, uterusexternushasbeenunknownuntilnow.Wefoundthatit
PME-ALE separation 0.08. Chelicerae length 1.02 with two islocatedbetweentheanteriormedianplateoftheAFGand
(tseleitghhtloynseppraormaatregd)in(Fi(gc.on7tAi)g.uoSuts)ernaunmd 1t.w0o3 loonng,re0t.r8o7mawrigdien, theVaproisatbeilriitoyr.—plaWteeohfavtheeePxaFmGin(eFdigst.he3fAe,maBl,e4gBe)n.italiaoftwo
morewidenedbetweencoxae2and3,sternumwithcuticular additional females from Neuquen and Rio Negro provinces i
projectionstowardcoxae(Fig. IF).Spination:LegIII:Tibiap (Argentina), one ofthem collected togetherwith two males.
0-1-0, r0-1-0, Vplap;metatarsusp0-1-1, r lap(displacedto The tip ofthe median rod and the size ofthe gland region
dorsal),v2ap.LegIV:Tibiarlap,v2ap;metatarsusp1-1-1,r differ in both specimens, but the morphology of the other j
w0-i1t-h1,abVouplt-ptle-n2.cutPiaclupl:arTalrosbuessdanpdi,twpo1,rovun2adpe.dTraercseapltoorrsgoann mplaaytesbedoselisghntoltyvvaarriya.blHeo,wmeavekri,ngretlhaetiivmempeodsiiatitoensdeotfertmhienpaltaitoens j
,
IZQUIERDO&LABARQUE—FEMALEGENITALIAOFORSOLOBUSPUCARA 513
E.FDiogrusraelss1hiAe-ldF.o—foOprissoltohhoussompau;caFr.aS(tMerAnCumN.-AScral1e61b2a0r)s.:FAe-mCale=.1A.mmH,abiDt,usEd=ors0a.l5;mBm.,HaFbi=tus0.2ve5ntmrmal.;C.Habituslateral;D.Eyesanterior;
Figures2A,B.—Orsolohuspucara(MACN-Ar 16567).Malehabitus.A. Dorsal;B.Ventral. Scalebars: 1 mm.
—
Figures3A-D.—Orsolohuspucani(MACN-Ar 10873).Genitalia.A-C.Femalevulva.A.Dorsalview;B.Lateralview,asteriskonthetwo
sclerotizedplates;C.Anteriormedianrod,arrowheadstotheglandducts,asteriskonrestofdigestedmuscles.D.Malepalp,leftprolateralview.
Abbreviations;b=booklung,pp=posteriorplate,E=embolus,ef=epigastricfurrow,mp=medianplate,mr=anteriormedianrod,PR=
posteriorreceptaculum,SA=spine-shapedapophysis,SD=spermaticduct,t=trachealtrunk,ue=uterusexternus.Scalebars;A=0.25mm,
B,C= 0.1 mm,D = 0.5mm.
of the species difficult. For correct determination, it is embolusseemsto originate froma fold ofa striated laminar
necessarytodissectanddigestthegenitaliaandthenobserve membrane (EM Eigs. 6B, C). Spine-shaped apophysis (SA,
thepreparation fromseveral pointsofview. Figs. 6B,C)closetothedorsallobe(DL,Figs. 6C,D).Dorsal
Male. DescribedbyForster&Platnick(1985).Weprovide subterminal lobe spine-shaped, ventral subterminal lobe
anadditionaldescriptionofthepalpofonemale(MACN-Ar slightlyflattened(DSL&VS—L, Fig. 6A).
16567)collectedtogetherwithseveralfemales. Internalcourse Other material examined. ARGENTINA: Neuquen Pro-
ofthespermaticduct(SD)asin Fig. 3D. Embolus(E)short, vince: Cerro Bayo, 1304 m, 40.74796°S, 71.59779°W, March
with a wide aperture at the tip (Fig. 6B). The base of the 2005, V. Werenkraut, pitfall traps (cod. M3S5M05), 2males
—
IZQUIERDO&LABARQUE—FEMALEGENITALIAOFORSOLOBUSPUCARA 515
Figures4A-E. Orsolohuspucani(MACN-Ar10873).Internalfemalegenitalia.A.Entiredorsalview;B.Anteriorportioninanteriorview;
C.Anteriormedianrod;D.Tipofthemedianrodshowingthepointsofmuscleattachmentsandrestofdigestedmuscles;E.Glandductsonthe
baseofthemedianrod.Abbreviations:AEG = anteriorportionofthefemalegenitalia, B = booklung,pp = posteriorplate,EC = external
cuticle,fa = flattenedapodemes, IF? = interpulmonary fold?, mp = median plate, mr = median rod, PEG = posteriorportion offemale
genitalia,PR=posteriorreceptaculum,T= trachealtrunk,ue= uterusexternus.Scalebars:A=0.5mm, B=0.2mm,C=0.05mm,D =
0.01mm,E=0.01 mm.
surFfiagcueressho5wiAn,gBth.—epOorrseosloohfusthpeiigccliarncid(duMcAtsC.NA-bAbrrev1i0a8t7i3o)n.s:PoBsSter=iobrasreecoefpttahecuglluamn.dAd.uctE,xtDerCna=lsduirstfaalcecasphoowfitnhgegtlheangdldauncdt,duPct=s;pBo.resI.ntSecranalle
bars:0.02mm.
516 THEJOURNALOFARACHNOLOGY I
vieFwi;guCr.esDi6ttAo-iDn.p—roOlratseorlaolbuvsiepwu;caDr.aD(itMtAoCrNet-rAolrate1r6a5l67v)i.ewL.efAtbbmraelveiaptailopn.s:A.DRLet=roldaotresraalllvoibeew,;DBS.LTi=pdoofrtshaelcsoupbutleartmoirnyalbulolbbe,inEdo=rsaelm-baopliucsa,l
LM = laminarmembrane,SA-spine-apophysis,VSL=ventralsubterminallobe.Scalebars:A= 0.5mm,B =0.05mm,C,D=0.1mm.
(MACN-Ar 19559);samedataJanuary2006(cod.M3S5E06), Chile and with pitfall traps in Neuquen and Rio Negro
L1opfeezm,ale15(02MmA,CN4-1.A0r994819°5S6,0);71.R5i5o80N1e°gWr,oMParrocvihnce2:006C,errVo. Provinces(Argentina). j;,
Werenkraut, pitfall traps (cod. M1S8M06), 1 female DISCUSSION — '
(MACN-Ar 19558). CHILE; Region IX, Cautin Province: Female genitalia, functional morphology. The peculiar '
Huerquehue National Park, Laguna Toro, in Nothofagus morphology of the anterior female genitalia appears to be
(Nothofagaceae)-ylrat/cr/r/a (Araucariaceae)-C/7M5q'ttra (Poa- adapted for muscle attachment. The places for muscle
ceae) forest, 995m, 39°08T8.7"S, 71°42'30.9"W, 7 February attachment seem to be restricted to the tip of the median -
21060152,0),Mv.ouRcahmerirceozdes&ARF.AMLRa0b0ar1q0u2e,5;1safmeemadlaeta(1MmAaClNe-aAnrd rofodthaendpoisttsevreinotrrapllapter.ojFeoctrisotnesra&ndPltaotntihcekfl(a1t9t8e5n)edhaavpeodneomteesd .
2immature(MACN-Ar 16568); 1 female(MACN-Ar 16570) that the lumen of the median rod of some species of
vouchercodeARAMR001026, preparationcode MAI-137; 1 Orsolobidae is sometimes heavily charged with sperm. The
female (MACN-Ar 10873), voucher code ARAMR000999, presenceofdifferentglandtypesintheanteriorandposterior >
preparationcodesMAI-99, 124, 138-140; 1 maleand 1 female portions ofthe genitalia has been taken as indicative oftwo E
(MACN-Ar 16567), male voucher code ARAMR000972, functionally different sites for sperm storage in the dysderid i|
preparation codes MAI-58, 69, female voucher code Dysderaerythrina(Walckenaer1802)(Uhl2000).Theseglands
ARAMR000971, preparation codes MAI-23, 63-68, 78; 1 would produce secretions generating different conditions of i|
male (MACN-Ar 16571), voucher code ARAMR001021, spermstorage,althoughothersecretionsmightbetransferred
preparation code MAI-128; Villarica Natl. Park, sector bythemaletogetherwith thespermatozoa(Burger&Kropf t
Quetrupillen, in forest of Araucaria, Nothofagus and Chus- 2007).Thepresenceofglandductsintheanteriormedianrod 1
qmuaelae,a1n2d803m,im3m9a°2t7u'r4e2.1("MS,AC7N1°-5A0'r441.625"6W9,),8MF.ebRraumairrye2z00&5,F.1 aofndOrtshoelreofboursepaucdaoruablseugfguenscttsiosn:omaettsatcohrmaegnetffuonrctmiuosnclaesswealnld, *§
LabDiasrtqruieb.ution,—PreviouslyknownfromArgentina,Neuquen sthpeerfmemsatloreaggee.niStaolmiea ocfoutlhdembuescilmepsliicnattehdeainntemreiocrhapnoirtsimosnooff jf
Province, here reported for Rio Negro Province and from sexualselection,asoccursinotherfamilies. Forexample,the
Chile,Cautin Prov—ince(Region IX). musclesM3,M4andM7canmovesomeplates,whichleadsto jii'v
Natural history. Orsolobus pucara was captured beating theclosingoftheuterusexternus in TriaerisstenaspisSimon
othnevCehigiestiapttieoanbianmabNoootsho(fRaagmuisraenzd&AraLuacbaarriqaufeorepsetr,s.esopbesc.i)alliyn m1u89s1c;lewhMe3resaesemisn tBoriegnnaobllieafreemcaolnedsittao(mCohvicekearibnuglge19c5l1o)sethtoe |
IZQUIERDO&LABARQUE—FEMALEGENITALIAOFORSOLOBUSPUCARA 517
Figures7A-E.—Orsolobiispucara(MACN-Ar16567).Female.Leftchelicera.A.Posteriorview,blackarrowheadstotheretromarginalteeth,
whitearrowheadstothepromarginalteeth.B-E.LeftlegIstructures.B.Tarsalclawsindorsal-apicalview,asteriskonthedistaltarsalorgan;C.
Clawsinretrolateralview;D.Tarsalorgan;E. Metatarsaltrichobothrialsocket.Scalebars;A-C = 0.1 mm,D,E =0.01 mm.
thegenital opening, whichmay lead to theejection ofsperm osmoregulatory processes is still unclear. Similargland ducts
(Burger2009, underOpopaearecondita). havebeenobservedintheoonopid UnicorncatleyiPlatnick&
In species of the genera Myrmopopaea, Grymeus and Brescovit 1995(M.A. I—zquierdo pers. obs.).
Lionneta, the surface of the posterior receptaculum is Phylogeneticcontext. IfthefemalegenitaliaofallDysder-
pervaded with papillae that resemble those present in the oidea are compared in an hypothetical evolutionary context
genitalstructuresofwatermites(Burger2010).Likewise,these with the hope of identifying homolog structures (Fig. 8),
papillae are present in the segestriid genus Ariadna (P. Segestriidae and almost all the Dysderidae fit well with the
Michalik pers. comm.). Apparently, the papillae might have notion of a typical Dysderoidea (that is, well delimited
afunctioninosmoregulatoryprocessesandcouldbeinvolved anterior and posterior receptacles), while the Orsolobidae
insperm activation (Burger2010). However, theglandducts and Oonopidae have complex anterior female genitalia with
ontheposteriorreceptaculumofOrsoiohuspucaraareslightly bizarresclerotizedelements.However,itisstillpossibletofind
differentcomparedwiththesespecies,henceitsinvolvementin similarstructuresandinfercommonmechanisms.Themedian
518 THEJOURNALOFARACHNOLOGY
1234567 1234567 4Anteriorreceptaculumsclerotization
DNAHyoansrpadpsdeasnrcpata-ebeaoreyltseheprinidbnaeargi 01l10O-1l00l10O000l100O00AMUnoytnronmoeonlpoaoppsspa.ceoarbsupl.o 111011000111011111100 HopsI sp.Aria--dHDnayarspdbaeocretaseeeanrbyletehprrigidinaa ”™HiSNAcmolebnriosgctulieozrueodstized
Orsolobuspucara 1011010Grymeussp. 1101110 -Osomolobussp.
Osomolobussp 1011010Orcheslinasp.1 1001010 -Orsolobuspucara
Opopaeafosuma lOOlllQOrcheslinasp.2 1001010 -Scaphiellahespera
SilhoueUellaloiicalula1-01110Scaphiellahespera 1001001 -Antoonopscorbulo
-Silhouettellaloricatula
-Opopaeafosuma
1Posteriorreceptaculum ....Absent Orcheslinasp.1
Nopssp.Ariadnaboesenbergi m^mAPrmebsiegnutous “IEOGrrcyhmeesluisnasps.p.2
HDayrspdaecrtaeearyltehpriidnaa LW’lyornmnoetpaopspa.easp.
OOsrsoomloolboubsupsucspa.ra 5Spermdumping MBAPbrseseenntt(Reported)
SAcnatpohoinelolpashceosrpbeurloa Nopssp Ambiguous
OSiplohopuaetetaellfaolsourmicaatula I AriadHnaarpbaocetseeanbleeprigdia
Orcheslinasp.1 I Dysderaerythrina
IEI OGrrcyhmeesluisnasps.p.2 _ OOsrsoomloolboubsupsucspa.ra
MLlyornmnoeptaopspa.easp. 1——SAcnatpohoinelolpashceosrpbeurloa
Silhouettellaloricatula
2Platesfusedtoposteriorreceptaculum Opopaeafosuma
Nopssp. Absent ““7I OOrrcchheessttiinnaasspp..21
_-IAriaHdDnayarspdbaeocretaseeeanrbylteehprrigidinaammmIAPrmebsiegnutous —irI MLGlyrorynmmnoeeptuaospsspap.e.asp.
71 OOsrsoomloolboubsupsucspa.ra 6Lockinguterusexternus Absent
']II11 ijSASOicnplatohpoophuOOoiaerrnetecclothhaelpeelaslssfahtlociielsoonnsruraapbmieucssaarlpptoa..ul21a --Nopssp.IAri———aDOdHOysnarsoarsdmpobeoalorcloetaboseuebeasnurbylpseteuhprcsrigpaid.irnaaa MMAPrmebsiegnuto(uIsnferred)
I' EMLGlyrorynmmnoeeptuaospsspap.e.asp. ———ScSAianlpthhoouioeentltoleplaslahceloosrrpbieucarltoaula
— —Opopaeafosuma
3AnteNroiporsrspe.cAerpiatdancaublouemsegnlbaenrgdis MSSAAPbrmsebseiengntutous -JII1I MOOLGrrlyrccoryhhnmmeenosseeptttuiiaosnnpsaasapp.sse.ppa..s21p.
--Harpactealepida 7Entelegynecondition
—DOOysrssodomelorolbaoubesruypstuhcsrpai.rnaa , Nopssp.Ariadnaboesenbergi mmP•Arbesseenntt(Haplogyne)
———SAOScinplatohpoophuoiaenetelotaleplaslfahocelsoosrurpbmieucaarltoaula II ' I1 _7I1 HDOOaysrrssopdomaelocroltbaoeubeasruylpsteuhpcsripaid.rnaaa Ambiguous
—^1,I1 MOOLGrrlyrccoryhhnmmeenoessettptuiiaosnnpsaaspap.sseppa..s21p. 7'IIII1—Ii,SAOScinplatohpoophOOuGoiaerrnretecclotyhhalepmeelasesslfahttuocieilssoonnsruraapsbimpeucssa.arlpptoa..ul21a
I J1 MLlyornmnoeptaopspa.easp. 8
—
Figure8. Datamatrix(upperleftcorner)andoptimizationforsevengenitalandsexualbehaviorcharacters.
rod and the lateral apodemes of the posterior plate in figs. lA,2A,8H-0;Uhl2000).AsForsterandPlatnick(1985)
Orsolobus are very similar to other species of Dysderoidea mention, there is a tendency for the storage function ofthe
(compareFig. 3Awithfig.2inBurger&Kropf2007andfig.3 anterior genitalia to become reduced as the posterior
in Burgeretal.2006).Thepresenceofglandductsandsperm receptaculum becomes larger. The absence ofgland ducts in
insidetheanteriormedianrodsuggeststhattheseandsimilar the oonopids analyzed here (Character 3, Fig. 8) and the
structures in the Oonopidae and other Orsolobidae are sclerotization of the anterior receptaculum (Character 4,
homologous with the membranous anterior receptaculum Fig. 8) seem to indicate a switch in the function of the
found in Dysderidae and Segestriidae (see Grismado 2008 anterior receptacle, from sperm storage to attachment of
IZQUIERDO&LABARQUE—FEMALEGENITALIAOFORSOLOBUSPUCARA 519
muscles involved in copulatory and post-copulatory mecha- EAR-0228699 and for a Planetary Biodiversity Inventory of
nisms. Gland ducts in the anterior female genitalia have thespiderfamilyOonopidae,grant0613754),ANPCyT(grant
recently been observed in undescribed oonopids from the PICT-2007-01393) and CONICET (grant PIP 112-200801-
mollesspinygroup(C.J.Grismadopers.comm.),Heteroonops 03209anddoctoralfellowshipsforM.A.I.andF.M.L.).TNT
Dalmas 1916(N.I. Platnick&N. Duperrepers. comm.), and was made available thanks to the sponsorship of the Willi
in Unicorncatleyi(M.A. Izquierdo& Rubiounpubl. data). HennigSociety.
Allthe Dysderoideaincluded in thematrixexceptAriadna LITERATURECITED
boesenbergiKeyserling 1877andScaphiellahesperaChamber-
lin 1924 have a mechanism of uterus externus locking Baehr,B.C.&H.M.Smith.2008.ThreenewspeciesoftheAustralian
p(flroCrechoksamiernnaggcecetttemirenogcf6h,ianmtnFuoiisgisc.tlmde8u)srisitnahgpanotdcsoswpisuobcllulealetdriobotenpicrza(eauBtvsuieoernngteortfohefettahtlse.hpe2ec0ro0ma6mna)btt.eiorTnziheoodear BreogRDsreeecssoncovcuoilrsrtoid,bpsLitodiAsoo.dfnsDoptl.aiho,ndehedirWtLsese.csAgotP.ellenoarugtBnysneirAoctHufkoisncttc&rkweamlolalBinonra,oeenlswocRMhos.iuvpiiesstcOeiteu(t(sAmArro&aa2fnn4ete:ahAa3ee.e2;A,5b.-rO3aDrz3yisL6slio.dsileeao.nrboisid2pda0iee0d)a4e.,.r
receptaculum (or part of it) and additional plates, both Orsolobidae). RevistaIbericadeAracnologia9:249-257.
servingas attachments for those muscles. When the muscles Burger, M.2007. Spermdumpinginahaplogynespider.Journalof
contract,theplatescontacteachotherandlocktheuterus(for Zoology273:74-81.
detailed morphology see Uhl 2000; Burger & Kropf 2007; Burger, M. 2009. Female genitalia of goblin spiders (Arachnida:
Fannes&Jocque2008;Burger2009,2010; Burgeretal.2003, Araneae: Oonopidae); a morphological study with functional
2006, 2010). The absence of sclerotization in the female implications.InvertebrateBiology 128:340-358.
genitalia of the segestriid Ariadna boesenbergi suggests that Burger,M.2010.Complexfemalegenitaliaindicatespermdumping
thismechanismisnotpresentinthis speciesand probablyin in armored goblin spiders (Arachnida, Araneae, Oonopidae).
tEuhnseicdaipwrhheiocetilileoanafhlaemsisplepyre.armTishfeclounxasbiissnetenntctheewoigftenhiltotachlekiai,ndgeavemlceoocnphfmaiengnuitrsamotifoiann BurZ(sgtAoerrorua,lccothugnMryi.ed,s1a1,i3Wn:A.d1ri9ac-anN3tee2ea.nset,wcriOygopntoi&cpifdCea.mea,lKeGraocpmhfao.siocme2o00ri3pn.hihaCnpaolemo)pg.lyenJxeoursgnpeainldietroalsf
typicalfortheEntelegynae(Character7,Fig. 8).InE. hespera Morphology255:80-93.
there are two ducts: one of them connects the copulatory Burger,M.,W.Graber,P. Michalik&C. Kropf.2006.Silhouettella
opening with the posterior receptaculum and the other loricatula(Arachnida, Araneae,Oonopidae); A haplogynespider
connectstheposteriorreceptaculumwith the uterusexternus with complex female genitalia. Journal of Morphology 267;
(Burger 2009). This configuration suggests that the locking 663-667.
mechanism of the uterus externus is not necessary in this Burger,M.&C. Kropf.2007.Genitalmorphologyofthehaplogyne
species, since the males have no direct contact with this spider Harpactealepkla (Arachnida, Araneae, Dysderidae). Zoo-
srterpuocrttuerdefdourrianngothceorpulgar.ouTphoeflgoacmkaisngommoercphhainniesmspehcaiessb(eneont Burmgoerr,phoMl.o,gyM.126I:z4q5u-i5e2r.do & P. Carrera. 2010. Female genital
morphology and mating behavior of Orchestina (Arachnida;
analyzed here; see Burger et al. 2006), and it is probably Araneae:Oonopidae).Zoology 113:100-109.
pIrzeqsueinetrdion&thLeagbeanruqsueOrpcerhse.stoibnsa.).as well (Burger et al. 2010; EbeFrehmaarlde,CWh.oGic.e.19P9r6i.ncFeetmoanlUeniCvoenrtsriotly:PrSeesxsu,alPriSnelceecttoino,nNbeywCJreyrpsteiyc.
Spermdumpingisacommonmeansofcrypticfemalechoice Fannes,W.&R.Jocque.2008. UltrastructureofAntoonops,anew,
bywhichthefemalesdiscardspermfromcurrentorprevious ant-mimicking genus ofafrotropical Oonopidae (Araneae) with
matings(Eberhard 1996). InDysderoideaspermdumpinghas complex internal genitalia. American Museum Novitates
been reported only in Silhouettella loricatula (Roewer 1942) 3614:1-30.
(Burger2007;Burgeretal.2006).However,thismechanismof Forster,R.R.&N.I. Platnick. 1985.Areviewoftheaustralspiders
cryptic female choice has been suggested for other gama- family Orsolobidae (Arachnida, Araneae), with notes on the
somorphine oonopids of the genera Opopaea and Xyphinus superfamily Dysderoidea. Bulletin ofthe American Museum of
SanidmoMny1r8m9o3,poGpaameaaso(mBourrpgehraeKtarasl.ch2010838;1,BGurrygmeerus2,010L)i.onTnheitsa GolNfooabrtoufprfhay,llPoH.gAie.sn,teotJri.ySc.a1Fn8aa1rl:ry1is-si2s&2.9.CKl.aCd.istNiicxson2.4:2707048-.78T6N.T,afreeprogram
behavior seems possible only with thecombined presence of Grismado, C.J. 2008. A taxonomic revision of the spider genus
sclerotizedstructuresandmusclesinthefemalegenitalia. Ariadna Audouin, 1826 in Argentina and Chile, with the
ACKNOWLEDGMENTS descriptionoffivenewspecies(Arachnida,Araneae,Segestriidae).
Zoosystema30:333-360.
Wesincerely thank Barbara Baehr, Barbara Knoflach and Griswold,C.E.&N.I.Platnick. 1987.OnthefirstAfricanspidersof
Matthias Burger for critical corrections on the manuscript. thefamilyOrsolobidae(Araneae,Dysderoidea). American Muse-
Martin Ramirez, Cristian Grismado and Norman Platnick umNovitates2892:1-14.
providedusefulcommentsonearlyversionsofthepaper.We Huber, B.A. 2002. Functional morphology ofthe genitalia in the
thanktheArgentinianAdministraciondeParquesNacionales spiderSpermophorasenoculata(Pholcidae,Araneae).Zoologischer
faonrdtRheesaeurtvoariNzaattuironaltoOtwaomrekndiin,twhheeCraeliOlrecghueastNiantaiosnpaelciPmaernks HubAgeenrnzi,etiaBgl.ieAar.,i2n4A1t.:wD1o.05nB-re1ews1c6s.opviidter&s(CA.raRnheeaiem,s.Ph2o0l0c5i.daEex)a,gwgietrhatceodmfmeemnatlse
wtheereScEoMllelcatebdo.raFtoarbyi.anThTeriscatruidcyowoafsfefriendanteccehdnibcyalgrsaunptpsorftroamt coonevogleuntiitoanl.IenvsoelcuttiSoynstebmyaticfsem&aleEvoclhuotiicoen3v6e:r2s8u5s-29a2n.tagonistic
thefollowingorganizations:NationalScienceFoundation(for Platnick,N.I.,J.A.Coddington,R.R.Forster&C.E.Griswold. 1991.
theAssemblingTheTreeofLife: Phylogenyofspiders,grant Spinneret morphology and the phylogeny ofhaplogyne spiders
520 THEJOURNALOFARACHNOLOGY
(Araneae, Araneomorphae). American Museum Novitates 3016: Ramirez, M.J. 2000. Respiratory system morphology and the
1-73. phylogeny of Haplogyne spiders (Araneae, Araneomorphae).
Platnick, N.I. & A.D. Brescovit. 1994. A new genus ofthe spider JournalofArachnology28:149-157.
familyOrsolobidae(Araneae,Dysderoidea)fromBrazil.American Uhl,G.2000.Twodistinctlydifferentspermstorageorgansinfemale
MuseumNovitates3112:1-6. Dysderaerythrina(Araneae, Dysderidae). ArthropodStructure&
Platnick, N.I. &N. Duperre. 2009.TheAmericangoblinspidersof Development29:163-169.
thenewgenus Escaphiella(Araneae,Oonopidae). Bulletinofthe
AmericanMuseumofNaturalHistory328:1-151.
Platnick,N.I.,C.J.Grismado&M.J.Ramirez.1999.Onthegeneraof
the spider subfamily Otiothopinae (Araneae, Palpimanidae).
AmericanMuseumNovitates3257:1-25. Manuscriptreceived31July2009, revised7August2010.