The Evolution of Prey-Wrapping Behaviour in Spiders Gilbert Barrantes A; William G

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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 The evolution of prey-wrapping behaviour in spiders Gilbert Barrantes a; William G. Eberhard a a Escuela de Biología, Universidad de Costa Rica, Ciudad Universitaria, Costa Rica

Online Publication Date: 01 January 2007 To cite this Article: Barrantes, Gilbert and Eberhard, William G. (2007) 'The evolution of prey-wrapping behaviour in spiders', Journal of Natural History, 41:25, 1631 - 1658 To link to this article: DOI: 10.1080/00222930701464364 URL: http://dx.doi.org/10.1080/00222930701464364

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Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 rus ‘oisre’o ‘ryrlig’wapn eaiu,i hc h pdrplsa pulls spider the which in behaviour, ancient. wrapping very ‘‘prey-rolling’’ be also or may ‘‘Rotisserie’’ wrapping groups. prey so 1989), al. et (Shear fossils ancient History Natural of Journal O:10.1080/00222930701464364 online DOI: 1464-5262 print/ISSN 0022-2933 ISSN Biologı de Escuela Eberhard, William Correspondence: in used also probably is silk silk post-immobilization gland aciniform wrap ampullate from pholcids serve while and (‘‘post- come clavipes of glands, forthcoming); Nephila to also al. acciniform feeding presumed et functions lacking can (Weng is or despite evidence prey These direct prey wrapping wrap little site, 1988). to is attack there use feeding Nitzsche although spiders glands, and the that 1975; silk exclusive, at Robinson The Mirick functions. not web 1974; and are Robinson the Knost 1969; wrapping and to al. et Rovner it from Robinson it 1971; fastening 1967; removing (Eberhard as it, so such wrapping’’) activities prey immobilization carrying other active facilitate restrain web, manageable to to compact, prey the prey: more immobilized form their already to wrap 1996). of and spiders (Foelix packages wrapping’’); prey which (‘‘immobilization wrap in to escape of contexts lines cannot feature two silk they such defining least use a at spiders is are many spinnerets and There abdominal 1989), from al. emerge et (Shear that spiders lines silk of production The Introduction construction egg-sac from evolved originally wrapping a at prey Keywords: Using convergently Scytodidae. that evolved and has propose Filistatidae IV in legs We time with first wrapping Agelenidae. times. behaviour. the prey and for that and four and Tengellidae, described propose linking prey is Theraphosidae, IV we least synapomorphy the prey spiders, in legs a wrap over of occurs both to be stands phylogeny it IV with recent may spider ancient; legs silk) legs, the very of both viscid which use apparently The of in with is movement other, place, covered The in simultaneous (often turns a Synotaxidae. line and with tense publications Nesticidae, prey described. previous a are Theridiidae, the combining holds techniques to by prey-wrapping spider it prey new Two the on applies families. attacks which 16 in in in spiders species One, by 31 of use observations silk new of with evolution the traced We Abstract ( Biologı de Escuela EBERHARD G. WILLIAM & BARRANTES GILBERT spiders in behaviour prey-wrapping of evolution The [email protected] cetd2 a 2007) May 21 Accepted rywapn eaiu a enue seiec fterltosisbtensome between relationships the of evidence as used been has behaviour Prey-wrapping takbhvor hlgn,sies rpigsilk wrapping spiders, phylogeny, behaviour, Attack a nvria eCsaRc,Cua nvriai,CsaRica Costa Universitaria, Ciudad Rica, Costa de Universidad á, W brad nulse) osbeaiiomgadsiosocri very in occur spigots gland aciniform Possible unpublished). Eberhard, (W. 07 12–8:1631–1658 41(25–28): 2007; , # 07Tyo Francis & Taylor 2007 a nvria eCsaRc,Cua nvriai,CsaRc.Email: Rica. Costa Universitaria, Ciudad Rica, Costa de Universidad á, Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 ou nvdoiae eeoitdfo rwnsbsdo hs images. in these not on were based that drawings Details from visibility. omitted to were according images varied video often in species given focus a in behaviour lines for of them. line’’ on ‘‘sticky liquid phrase then This the of hours, microscope. use balls several We dissecting visible see. to a to easily min under easy 40 with lines very for balls the towels hydroscopic observing wet made quickly treatment with and lined chamber chamber the ‘‘humid’’ opening small a in prey e ftesie.Uls oe tews,alsieswr auefmls l en are means All females. mature were spiders all otherwise, noted by Unless followed spider. the of web oahr otesie’ e,s twsfrtetnldi i fcielmsl rmteweb the from silk cribellum of bit failed a prey in the entangled cases first few spider; was a the it In a so near web. web, spider’s of dropped spider’s the were the to theraphosid on to DCR dropped adhere the species were to Sony for different species Prey and for other I. the used Table DCR-VX1000 for prey in prey Sony The listed lenses. using are close-up noted) recorded with (when were equipped cameras prey digital to TRV50 spiders of Responses in Methods occur well may details ‘‘new’’ previously. these studied of been to some have IV legs Therefore that of prey. species use the simultaneous other such versus to ‘‘new’’, details, alternate silk subtle as the wrapping some or details of spinnerets, apply occurrence different behavioural temporal the of some for positions fine checked the present the be as to the We cannot in before studies behaviour. and used or previous wrapping been most however, on but never of them alternate apparently superficial, details has place make often the recordings and IV study video are digital spinnerets legs Descriptions from the spider’s available prey. from resolution the the lines that of silk statement wrapping vicinity common pull the to than movements other 1967). 1974), prey, (Eberhard Knost wraps diguetids and and Rovner 1986), in evolved (Eberhard (Melchers scytodids and ctenids silk 1974), (Rovner vegetation 1988), (Valerio in wrapping drymusids groups, (Nitzsche ground the 1990). pisaurids several above Viscid 1974), in Opderbeck live that Knost convergently, and 2004). lycosids perhaps some (Kirchner others Agnarsson occurs, among Pholcidae including also family 1998; wrapping related prey (but al. distantly synotaxids Post-immobilization and the aggregate et theridiids, the in nesticids, by (Griswold convergently linking produced synapomorphy The below) presumably a 1998). silk, be al. see viscid to et with thought Griswold covered is lines line 1982; glands, the silk a (Eberhard on is wrapping araneids prey 1971), of the Olizarri of spinning and use subfamily by Robinson 1969; prey one Robinson the of 1933; to 1931, synapomorphy on (Peters it entangled wraps is and it spinnerets where its from silk of swath 1632 eddt s eaieylrepe htwr ifcl o h pdr oesml ie are sizes sample Some spider. thus the We for conditions. difficult were certain that under prey omitted large was relatively 1969; behaviour pattern, use were behaviour this al. to given we not a et study tended perform or this (Robinson could whether In species prey than 1980). a not the Lubin rather or 1974; of whether Knost determining group and in taxonomic Rovner interested 1971; and Olizarri size and the Robinson as such circumstances and rvossuishv hw hti oegop,sc as such groups, some in that shown have types studies different of Previous sizes sample the and visible, not were details some recordings some In wrapped recently placing by lines wrapping on material adhesive of balls for checked We ulse ecitoscnanfwdtiso h oeet ftesie’ eswieit while legs spider’s the of movements the of details few contain descriptions Published uucnahibernalis Kukulcania Lycosa .Brats&W .Eberhard G. W. & Barrantes G. ¡ rpigbhvorvre ohqaiaieyadqatttvl with quantitatively and qualitatively both varies behaviour wrapping , n tnaddeviation. standard one Hnz 82 n hntecielmsl a nage nthe in entangled was silk cribellum the then and 1842) (Hentz, Argiope , Cyrtophora , Nephila , Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 auefml)wsfdsvrllvn oh htwr rpe – mfo e.Between her. from cm 1–5 dropped were that moths living several fed was female) mature calidus vicina Cicadellids nymphs Membracid sloou reduncus body its Psalmopoeus moving by silk wrapping pulls Spider will preparation). IV in legs Barrantes, their G. with and terminated them Eberhard and released II. G. then Table initiated (W. some and spiders in elsewhere them of described snagged how summarized and they be concerning how species, are and details representative lines, sizes Further wrapping by sample follow. movements details and wrapping unusual behaviour particular of wrapping Descriptions of aspects Several Results Vouchers Ubick. Darrell Zoologı by de Museo defined) the clearly in not deposited are been limits have (generic identified tentatively and 1997); (Levi Spiders butterflies Moths, spiders. of species different Prey to presented prey of Types I. Table hibernalis and 1874) Levi]; bifurca were Allocyclosa we which species. in another behaviour particular to a on execute moved to we found was interested, species a once because small, a ete,wss bees wasps, Beetles, spider Araneid eaoeosflies Nematocerous uci flies Muscoid rspii flies Drosophilid ml clprt flies acalyptrate Small esdflies Sepsid Ants Roach Chrysopid Pseudomyrmex pdr eeietfe ytefloigseilss(nsur rces n references: and brackets) square (in specialists following the by identified were Spiders O .Cmrde 89 B pl] l theridiids, all Opell]; [B. 1899) P.-Cambridge, (O. ctdslongipes Scytodes etc,17 I Agnarsson]; [I. 1971 Gertsch, Uik2005); (Ubick Ochyrocera sp. b okr n odesof soldiers and Workers d MCo,18)and 1887) (McCook, zlaaffinis Azilia p B Huber]; [B. sp. hibernalis caaae tesselata Achaearanea Dubiaranea tesselata Achaearanea ehl clavipes Nephila ctdslongipes Scytodes hys intervalensis Chrysso hys cambridgei Chrysso tesselata Achaearanea geometricus longipes tesselata Achaearanea evexum eau gravidus Metabus caaae tesselata Achaearanea caaae tesselata Achaearanea Deinopis cambridgei Chrysso ua,14 Vlro1981); (Valerio 1844 Lucas, (Theraphosidae). eglaradiata Tengella , , Tortolena uuknahibernalis Kukulkania sp. b , , ctdslongipes Scytodes sp., ehl clavipes Nephila .P-abig,19 Lv 1980). (Levi 1893 P.-Cambridge, O. , Synotaxus vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution Pheidole , aoe sellata Kapogea , sp. Scytodes sloou reduncus Psalmopoeus , aclu calidus Gaucelmus , , , , , , evbslongipes Helvibis hlpnlavicina Philoponella zlaaffinis Azilia .tepidariorum A. nlsmsstudiosus Anelosimus loyls bifurca Allocyclosa nlsmsstudiosus Anelosimus Dubiaranea ecuemariana Leucauge Klznk,10)(of 1977); (Wolff 1909) (Kulczynski, nec ffv aigssin h pdr(a spider the sessions taping five of each In sp. sp., sp. c , , c etcie rufipes Nesticoides ao h nvria eCsaRica. Costa de Universidad the of á hrdo evexum Theridion igdmale Winged , Ochyrocera hlpnlavicina Philoponella , sloou reduncus Psalmopoeus , , , hlpnlavicina Philoponella evbslongipes Helvibis nlsmsstudiosus Anelosimus , ? p G Hormiga]; [G. sp. (?) , nlsmspacificus Anelosimus hsccu globosus Physocyclus Ochyrocera a , sp. , .pacificus A. Camponotus Chrosiothes Krc,18)and 1880) (Karsch, , , Synotaxus Tcaosi 81 H W. [H. 1881) (Taczanowski, Ochyrocera Synotaxus , hsccu globosus Physocyclus sp. , , , p nr. sp. eglaradiata Tengella Kapogea hys intervalensis Chrysso , sp. Ochyrocera sp., , sp. .pacificus A. d , Trigona p and sp. hsccu globosus Physocyclus hrdu evexum Theridium porteri sp., Melpomene (Taczanowski, , sp. Latrodectus sp. , Kukulkania Kapogea , , Kukulkania Philoponella Theridium zlaaffinis Azilia Tortolena Gaucelmus , Helvibis 1633 was sp. sp. , Linyphiidae Scytodidae Uloboridae Agelenidae Deinopidae Tengellidae Ochyroceratidae Theraphosidae Pholcidae Filistatidae Theridiidae Taxon spiders. of species different of behavior Wrapping II. Table Dubiaranea ctdslongipes Scytodes hlpnlavicina Philoponella Tortolena nlsmsstudiosus Anelosimus pacificus Anelosimus Deinopis radiata Tengella Ochyrocera sloou reduncus Psalmopoeus hsccu globosus Physocyclus hibernalis Kukulcania caaae tesselata Achaearanea iorum tepidar- Achaearanea p Alt ? ? sp. p oN No – – – – No No sp. sp. p oA least At No sp. h Immobilization wrapping Yes/no edmNo Seldom Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 No e oAt–0(1 9,3)Yes 30) 196, (211 0 – Alt No Yes e Sim – Yes e l 84 )?YsYes Yes ? 2) (804, 0 – Alt ? Yes e Sim Sim ? ? Yes Yes No oN No – – – – No No oN No – – – – No No f i g a Spinnerets ihlegs with touched close e l 52 )Ys?Ys2 lgtyN I I Yes III II, No Slightly 2 2, Yes ? Yes No 1) (512, No 0 – – Alt – Yes Alt Yes IV o8 – – – 3 8, No – – – – – – b s Simultaneous At s of use (Alt) Sim alternate Sm or (Sim) esIV legs l (e? Yes ? ?(Yes?) – – Alt + + + + k Alt l e 0 – – – – 1 10, – – – – Yes Alt l e 6(1,1 e e ,1???Yes ? ? ? 1 1, Yes Yes Yes Yes No ? 6) (446, 53 1) (113, 26 Yes Yes Alt Alt a t Simultaneous rqetin frequent s flegs of use wrapping Vmore IV early No ––?No u n.movements, (no. simultaneous 0(5,9 e e e 0 e oIII p I–III, No Yes 5 10, Yes Yes Yes 9) (351, 20 movements flg IV legs of o prey) no. %of rywith prey esIV legs Touch m c wrapping iein line Sticky Yes early n,o d d d d l observations oevideo some tleast At Yes Yes e ,3Ys––– – – Yes Yes 3 8, Yes Yes o1 ? ? ? ? 1 1, No e e f . N spiders) . . . . . 20, . . ,4Ys?III ? Yes 4 4, 0 e – – – Yes 2 20, 0 e ,IIIYes – I–III p, – ? – Yes 2 30, Yes 1 30, (prey, 10 20 50, 50, . e oI,III II, No Yes 5 spinnerets wrapping lgty?I,IIYes III II, ? Slightly Spread e e I I Yes III II, Yes Yes as esI as IV legs applied spread Swath with managing ap(p) palp esor Legs sdin used prey j p r v iea leg as side Abdomen Vgrasps IV wn to swung Yes Yes Yes line

–

1634 .Brats&W .Eberhard G. W. & Barrantes G. al I Continued II. Table Araneidae spp. Nephilidae Synotaxidae Tetragnathidae Nesticidae Taxon porteri Chrosiothes loyls bifurca Allocyclosa ehl clavipes Nephila hys cambridgei Chrysso aoe sellata Kapogea affinis Azilia sisyphoides Tidarren rufipes Nesticodes Synotaxus ecuemariana Leucauge calidus Gaucelmus rip argentata Argiope hrdo evexum Theridion hrnii reimoseri Phoroncidia geometricus Latrodectus longicauda Helvibis intervales Chrysso ddd p or sp. p nr. sp. ww ? No (?) Immobilization wrapping Yes/no Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 e Sim ? Yes e oAt–0(5 )N Yes No 2) (45, 0 – Alt No Yes e Sim – Yes e Sim ? Yes No Yes e No Yes e oAt–0(7 )??Ys3 e ,I III I, p, ? Yes 1 3, Yes ? ? 2) (87, 0 – Alt No Yes e No Yes e Sim ? Yes e Sim Sim Yes Close Sim – Yes – Yes Yes oYsn l Yes – – Alt Yes/no No o?At–0(4,3 o?Ys4 ,I,IIYes III II, I, ? ? 3 4, Yes ? No 3) (241, 0 – Alt ? No k xx a Spinnerets ihlegs with touched IV oAt–0(9,4 Yes 4) (297, 0 – Alt Sim No No nn b ff s aa Simultaneous At s of use (Alt) Sim Sim Sim Sim alternate Sm or (Sim) esIV legs ll + + + + + + + + + + + + l e Yes ? – Yes Alt l No Alt l e 8(4,2 oYsYs4 e oI,II– III II, No Yes 2 4, Yes Yes No 2) (246, 38 Yes Alt l e 0(7,5 o?Ys5 e e ,III I, Yes Yes 3 5, Yes ? No 5) (478, 40 Yes Alt l No ? – ? Alt l –0 5– 5–10, 5– 5–10, – – – – – – – – – – Alt Alt l e 6(07 )L; 8) (1087, 36 Probably No Yes 2) (189, 31 4) (329, 35 Alt Yes Yes Alt Alt l e 6(6,3 No 3) (560, 26 Yes Alt l Yes – – – Alt a Simultaneous rqetin frequent s flegs of use wrapping Vmore IV early x n.movements, (no. 6(9 )S 3) (79, 16 simultaneous 3(4,2 oYsYs2 lgtyN I–III No Slightly 1 2, Yes Yes No 2) (748, 43 movements flg IV legs of o prey) no. %of oo rywith prey esIV legs Touch no oYes No aaa gg ccc bb zz c wrapping iein line Sticky Yes Yes early e Yes Yes oYs1,2YsN –I Yes I–III No Yes Yes 2 12, No Yes No mm eee hh pp cc w tt observations oevideo some tleast At Yes No e ,3??–– only? – Late Yes 4 8, 4 9, ? Yes Yes ? 3 3, Yes Yes Yes f e . . N spiders) . . 10 30, . 5,5???? 2,2–––– 0 e e ,I,IIYes III II, p, Yes Yes 5 30, 0 Slightly 3 30, (prey, 10 10 20, . 7Yes spinnerets wrapping Spread as –––– –––– ??p dd qq bbb ii esI as IV legs applied spread Swath Yes No with e I I Yes III II, Yes III Yes rr jj managing ap(p) palp uu esor Legs I III II, sdin used ,III I, prey I I ? III II, , ee yy ss y kk vv iea leg as side Abdomen Vgrasps IV wn to swung Slightly Yes line Yes Yes Yes Yes

vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution 1635 al I Continued II. Table lopls esII. legs palps, also rpigmvmnswr nrqet n h pdruulybtpe eoewapn;wapn eryawy curd u a eaieybrief. relatively was but occurred, always nearly wrapping wrapping; before prey bit usually spider the and infrequent, were movements wrapping ihaporaeageadfcs e Vddnttuhspinnerets. touch not did IV leg focus, and angle appropriate with e prahd vni ae ihsmlaeu e oeet htddnttuhtepe,teln ewe h espoal oce h ryo nearby or prey the touched w probably prey legs the the because between itself, line prey the the prey, not the if touch prey, not the did to that attached movements line leg moved. silk simultaneous prey a with the least cases at in touched Even often approached. leg leg the movements), simultaneous during occasionally aeyvisible. barely edmI hnbtn it. biting when I) seldom leosre nmcocp nlnscletdbt ietyfo h pdr n ntewapdprey. wrapped the on and spider, the from directly both collected lines on microscope in observed glue oehtsik Eehr ta.20) hssikns speual o ooooswt hto hrdisadnesticids. and theridiids of that with homologous not presumably is stickiness This 2006). al. et (Eberhard sticky somewhat ie ahrta wt flines. of swath a than rather lines ns fsvrltpso ryofrd eewapdwt tcysl ihsmlaeu oeet flg IV. legs of movements simultaneous with silk sticky with wrapped were offered, prey of types several of ants, hnte ntae wrapping. initiated they when h he e,adntmnpltdsubsequently. manipulated not and web, sheet the ryb ptigo t oeie rpigocre eoe n oeie fe h rywsbitten. was prey the after sometimes and before, occurred wrapping sometimes it; on spitting by prey lw n oedtiswr loesl bevdwt h ae eye. naked the with observed easily also were details some and slow, agru ryb iig nldn orccdliswihwr bu n-hr hi ieadkce nreial.Te edl prahdpoten approached readily They energetically. kicked and size their one-third about were which cicadellids four as including such biting, by prey dangerous a louiu.Tesie gi rk n eldu h ieo e a akt e etn iea n n ftehrzna ie n elcdi ihast a with it replaced and line, horizontal the of end one at behaviour site The resting web). her of entire to behaviour her back the constituted way to 180 which her identical turned line, on are sticky she line details horizontal the these place, up single, resting reeled (the her and prey reached broke the again she approached spider she The which unique. along also line was the up reeled and broke htapidsik silk. sticky applied that 17)as pcfe httelgI oeet of movements IV correct. leg is description the that specifies also (1971) hr a ocerpteni h eaiefeunyo iutnosadatraemovements. alternate and simultaneous of frequency relative the in pattern clear no was there hs pce ontwa ry si spsil htw i o iete h prpit ryt lctwapn attacks. wrapping elicit to prey appropriate the them give not did we that possible is it as prey, wrap not do species these flg Vtesinrt eeceryspread. clearly were spinnerets the IV legs of Eehr ta.2006). al. et (Eberhard ol o ueottepsiiiyta nemdaepstoswr isdi h ie record. video the in missed were positions intermediate that possibility the out rule not could asdna h pnees nohr h pgeo h aea oeeto h boe onie ihteapoc fteisltrllgI;tecoord the IV; leg ipsilateral directions. different the several of in approach sheet the the with to coincided prey abdomen the the anchored that of lines movement laying lateral by the followed of was apogee Wrapping the others. others in different in spinnerets; the near passed ee a iha es oeprey. some least at With esIIhl rydrn tlatls ato wrapping. of part last least at during prey held III Legs Solenopsis aa oeet o ai ob ueaotcnat u hr eemn rmsi hc tsee likely. seemed it which in frames many were there but contact, about sure be to rapid too Movements cc ns u undaa fe re otc ihteratro legs. anterior their with contact brief after away turned but ants, eylresik al ttevr is;i ae stages. later in first; very the at balls sticky large Very q h oriainwt e oeet a aibe nsm ae h boe oe lgtyt h ieadpseiry00–. fe h e IV leg the after s 0.07–0.1 posteriorly and side the to slightly moved abdomen the cases some in variable: was movements leg with coordination The l ie hce ne iscigmcocp akdsik al.Tebd fteflywapdpe a o opesd si yia nuloborids in typical is as compressed, not was prey wrapped fully the of body The balls. sticky lacked microscope dissecting a under checked Lines hh m loocsoal ae nwapn attacks. wrapping in later occasionally Also w rmrpd(0 rmsprs ie eodns(.D Bricen D. (R. recordings video s) per frames (500 rapid From b ra es eycoet h spinnerets. the to close very least at Or laigms iil nvdorcrig no otgost prey. to contiguous or on recordings video in visible mass Gleaming Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 s nfae fvdowt etfcslg Vapae otuhtesinrt,btnti h rmswt etveigangle. viewing best with frames the in not but spinnerets, the touch to appeared IV legs focus best with video of frames In ss tbgnigo taklg IadIIhl e hedt hc ryahrs ae r lc npe package. prey on place are later adheres, prey which to thread web hold III and II legs attack of beginning At jj .studo P. oeie u o always. not but Sometimes u n hnree nteln eidhr(tahdt h erysbtae ihatraemvmnso e esI.Alof All IV. legs her of movements alternate with substrate) nearby the to (attached her behind line the in reeled then and , k .longipes D. tlatatrtefrtmnt rs fwapn,wihwsntosre.Tedsrpino oisnadRobinson and Robinson of description The observed. not was which wrapping, of so or minute first the after least At Eehr 1981). (Eberhard ff ako otc o boueycran nsm eune tsee httetru lal i o oc,btwe but touch, not did clearly tarsus the that seemed it sequences some in certain: absolutely not contact of Lack eei lento ‘bccig’;snealwapn oeet eswwr ut lw essettheir suspect we slow, quite were saw we movements wrapping all since (‘‘bicycling’’); alternation in were c Usually. oo f ii h pdrddntbt oepe eoesatn owrap. to starting before prey some bite not did spider The ag ry()wr agrta osfy ml ry()were (S) prey small housefly; a than larger were (L) prey Large al nwapn pneeswr ee eni pedpsto;ltr uigatraemovements alternate during later, position; spread a in seen never were spinnerets wrapping in Early mm dd kk rbbysra,a es uigerywapn hnsmlaeu e Vmvmnsoccurred. movements IV leg simultaneous when wrapping early during least at spread, Probably hc,bih ht ieeegdfo pneesi ie recordings. video in spinnerets from emerged line white bright thick, A oeie lopalps. also Sometimes d rmvdoiae ny ie eenteaie nhmdchamber. humid in examined not were lines only; images video From i ako takwapn norosrain osntcnicnl eosrt that demonstrate convincingly not does observations our in wrapping attack of Lack o esnlcommunication). personal ˜o, gg y tlaterydrn rpig uigsmlaeu oeet flg IV legs of movements simultaneous during wrapping, during early least At x esIwr sdwt ag prey. large with used were I Legs h eyfrtmvmnswr iutnos u nters fwrapping of rest the in but simultaneous, were movements first very The ll takbhvordfee rmta fohrteiid nta h spider the that in theridiids other of that from differed behaviour Attack h pdr ncpiiyraiyatce oeaeszdadsomewhat and moderate-sized attacked readily captivity in Spiders qq ieysra,a es tedo rpigbout. wrapping a of end at least at spread, Widely v pdrgabdpe ihplsadlg IadII(but III and II legs and palps with prey grabbed Spider n oblso lepeet u eeteesa least at nevertheless but present, glue of balls No j rywr ezdi h hlcre uldthrough pulled chelicerae, the in seized were Prey bb o r e ege l (forthcoming). al. et Weng See pdr vie otc ihdneosprey dangerous with contact avoided Spiders hl rpigwt lent esI (and IV legs alternate with wrapping While g rpe nyatrfrtimblzn the immobilizing first after only Wrapped z htmcadmnlmvmnswere movements abdominal Rhythmic Drosophila tt rpiglnshdsmall had lines Wrapping rmosquitoes. or nn u Only nasnl ie frame video single a In e sdfetdwe the when deflected as pdrbhvorwas behaviour Spider ilydneosprey dangerous tially olwn nattack an following Pseudomyrmex nto a still was ination cyln.When line. icky t ie,because lines, Simultaneous p Occasionally rr w thick Two pp Viscous

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1636 .Brats&W .Eberhard G. W. & Barrantes G. eecletdatrbigwapd n lcdi ui hme o eoebigosre ne iscigmcocp,oehdn ie ihdrop with lines no had one microscope, dissecting a under it. observed on being droplet before small h 8 a for with chamber line humid a one in only placed had and others wrapped, being after collected were rya tsig nwbthread. web on swings it as prey aaiu,Hrda ot ia(lvto bu 0m n taot80mi aqeEtda nevls P rzl hymyb ifrn species. different be may they Brazil; SP, Intervales, Estadual Parque in m 800 about at and m) 20 about (elevation Rica Costa Heredia, Sarapiqui, rnia sucran(.A (F. uncertain is Araneidae u oeie lohl nwt n e Iwieuigteohrt oaeteprey. the rotate to other the using while II leg one with on held also sometimes but rpe first. wrapped asso iud(e ege l otcmn) u akdtelreblstpclo theridiids. of typical balls large the lacked but forthcoming), al. et Weng (see liquid of masses al I Continued II. Table yy esI n I eeol cainlyue ohnl prey. handle to used occasionally only were III and II Legs

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vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution 1637 Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 1638 eoelx(ecudntsetelxsl ntevdorecordings). video the have in must silk silk lax of the band see the not spinneret), could trailing (we lax the become for common (particularly attachments h rpiglnswr adudrtnin u hntesie turned spider the when but tension; under laid were lines wrapping the riigpseirsinrt rdcddfeetpten flns hnsuccessive When lines. of patterns different produced were spinnerets attachments posterior trailing hmbrl bv h usrt,sesughradmnt n ie(34 side one lifting to After abdomen b). her 1a, swung (Figure she times and more substrate, lines or the long one wrapping above her substrate initiated pressing barely the by spider them lines to the wrapping briefly initiated I, spinnerets apparently stage She posterior In substrate. the stages. to three them attached to up included chelicerea n hnt h te (20 other the to then and edn pnee a lss otesbtae h oa nl hog hc h spider the the which of through face 834 angle inferior total averaged the The of II 486.0 substrate. portion stage the lateral during to outer, closest turned the so was 1c), spinneret (Figure leading dabbed was she spinneret as leading slightly her with dabs between them she turned of As she both angle or one. spinneret mean 90 leading leading The the the substrate. spinneret either trailing became the dabbed the she to spinneret while Periodically trailing posteriorly, the 1c). (Figure directed from her raised was down turning was that spinneret it of posterior set so direction she leading I, later the the spinnerets. but turned, stage reversing her chelicerae, by of her from began in attachment silk prey spider last further the The held pulling wrapping. she thus II, continued stage and more and of around cephalothorax, part rotating her first place, video the in of During turned the centre spider the The of II. less e). stage frames or 1d, of (Figure scattered movements visible turning in bands the began the confirmed made band spider were illumination a the of of spinneret Attachments angles swing posterior substrate. favourable each the when each to recordings, During times from four pattern. silk to ‘‘zig-zag’’ one of spinnerets short posterior a her dabbed making sequences, wrapping led nhrceiea.Ol fe h tuge ftenwpe a usdda twas it as subsided moths had three prey to new begin. up the wrapping with did of prey chelicerae struggles captured spider’s the spider the of after The reoriented tip in Only then legs. posterior held and chelicerae. the front III, her near both leg in fell nearest with moth her already strike with and the to first prey, when strike body the case to toward one her her turned nearly In she with more I. abdomen, it face legs spider’s to both the grasped body with had her again reoriented struck she and then until prey IV the released substrate. legs briefly the except on legs placed to legs then all forward front were moving and or both swiftly legs one spider, pedipalps while Most the with even her, her of moth chelicerae. front toward the spider with in at it less The it struck pulled or then She hold side. more she simultaneously. was legs; her prey moth front the the to her When response struck lift legs. first or and front spider’s her prey, behind, The of the ground. (as both front, face the prey to in reached turn all prey to fell to falling was were rapidly a they attacks before and Nine whether responded accurately feeding. sometimes s), extremely was 0.06 responded spider as only the spider dropped quickly and was The moth subdued detail. new been each in session; had analysed each moth in spider previous the the to after fed were moths four and two pnee a 151 was spinneret eas hywr abdt h usrt ihdfeetfeunis h edn and leading the frequencies, different with substrate the to dabbed were they Because fe h atsig h pdrams lasdbe e pneesoc rtiea she as twice or once spinnerets her dabbed always almost spider the swing, last the After rpigbhvor( behaviour Wrapping then leg, front one with first struck sometimes spider the rear, her to fell prey a When ¡ 89 ¡ u 388.4 2 asi ih eune) h enagebtendb ihhrtrailing her with dabs between angle mean the sequences); eight in dabs (26 .Brats&W .Eberhard G. W. & Barrantes G. u hl odn h ry n h etatrstigi down. it setting after rest the and prey, the holding while ¡ , 265 180 N u u 5 2 asi ih eune) h pdrsadmnwstilted was abdomen spider’s The sequences). eight in dabs (27 pr a a atclrycmo o h edn spinneret), leading the for common particularly was (as apart ¡ 2 htocre hntesie led a te ryi her in prey other had already spider the when occurred that 12) 10 u , N 5 ¡ ) hs wnswr hnrpae nfu f10 of four in repeated then were swings these 3); 430 u ( N 5 1.Tesie unda vrg of average an turned spider The 21). . ¡ 180 18 u u , between N 5 12), Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 ob uldtgtyaanttetpadsdso h rypcae n obek stespider chelicerae. the her As break. with to repeatedly and bundle package, prey prey substrate the the the of to sides crushed attached and she top lines fed, the the against caused the tightly Lifting pulled cases away. be other to centimetres In several ones. walked also smaller she four least II. or four at stage three to captured after or one had feed long) with spider to cm prey 2 the started the when (ca of spider occurred moths surface wrapping large the III relatively and Stage forward. substrate two moved spinnerets. the she her to as of silk prey the of dabs over to bands forth and began the and posteriorly, then fastened back extended and laterally, She were times, abdomen spinnerets her four posterior swung to Both spider two 2a). the prey (Figure the lines zig-zag near more substrate spider lay the the to as spinnerets frequent both dabbed more was It time II. spinneret. leading stage same the of the from end at band the spinnerets attached neared frequently both more dabbing the that by appeared prey the to on pressed rpiglnseegn rmtetaln,rie pnee r iil;()wapn ie mrigfo the from emerging lines wrapping (e) visible; other are spinneret followed (arrow), (d) turns raised lines raised; visible. spider is trailing, are dotted spinneret the trailing spinneret the (c) by the lowered and substrate; from indicated lowered leading, the is emerging (position to spinneret leading spinnerets lines substrate the posterior while wrapping the right both the presses to toward spider abdomen pressed its the swinging (b) is s); 0.10 spinneret by posterior position right the as iue1 rpigbhvorof behaviour Wrapping 1. Figure atnn tloeyt h usrt.Telxsl htacmltdwe h nl between angle was the it, when spinneret around accumulated and trailing that surface the silk upper lax its with The when across attachments substrate. laid Later, successive the were prey. to lines the loosely substrate, touch it the not fastening on did were prey attachments presumably the successive attachments placed when other she only between lines laid the lines in and wrapped were prey the of obgnt ed h pdrgapdtebnl nhrceiea n itdi;sometimes it; lifted and chelicerae her in bundle the grasped spider the feed, to begin To spider the prey), the straddling while pause brief a after (sometimes III stage wrapping In hl h pdrhl h ryi e hlcredrn h is ato tg I h sides the II, stage of part first the during chelicerae her in prey the held spider the While sloou reduncus Psalmopoeus vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution Teahsde.()Sie it e boe oteright the to abdomen her tilts Spider (a) (Theraphosidae). . 180 u a oeie aebeen have sometimes may . 180 u apart, 1639 Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 ert ifrn prey. different to near tahetb ohsinrt;t tahetb riigsinrt;(,c tahet aeby made attachments c) (b, spinneret); trailing by attachment t, spinnerets; both by attachment oatc e ry iueo oatrgapn h rywt e hlcre hnthe when chelicerae, her with prey the wrapping. grasping began she after subsided, so ran substantially she or her as had I– minute of behind legs struggles left A both with were prey’s by prey. or prey prey one captured the new video) Previously securing as attack on chelicerae. forward, soon her to recorded rapidly and As moved sheet were anteriorly. spider the (eight extended against the III moth, legs attacks the front initiated contacted her She I with legs times. upper prey all the the on at to moved rushing sheet spider The the observations. of behavioural surface facilitating thus retreat, a lacked of that 1640 eglaradiata Tengella by lines wrapping of attachments of Patterns 2. Figure ertepe akg tiklns nmesrfrt re nwihatcmnswr ae a Zig-zag (a) made. were attachments which in order to refer (numbers lines) (thick a by package attachments prey the near he e ncpiiysoe httewapn eaiu of behaviour wrapping the that showed captivity in web sheet .reduncus P. .Brats&W .Eberhard G. W. & Barrantes G. P . (Tengellidae). reduncus ept h atta t otro pneesaemc hre.Ti web This shorter. much are spinnerets posterior its that fact the despite , nsaeI ro otrigaoeape l tahetb edn pnee;b, spinneret; leading by attachment (l, prey a above turning to prior I, stage in bevto ftremtr eae atrn oh na on moths capturing females mature three of Observation sloou reduncus Psalmopoeus .radiata T. a and (a) eglaradiata Tengella a ut iia to similar quite was .radiata T. b )o and on c) (b, nand on Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 abdi gis h he ma nl ewe asws94 was dabs between angle (mean sheet the against it dabbed vrtepe e ie Fgr b ) unn n rpigpe a eni nyoeof one passed only generally in on seen swath spiderlings was The prey instar abdomen. wrapping second and her of Turning of her c). attacks 2b, to tip 120 from (Figure attached the times about lines few was dabbed a of swath last prey the the swath had over that she a revealed where favourable, orientations sheet were their the visible; angles was lighting spinnerets when posterior Occasionally, prey. the wrapping). of h pdrdsperddw h unlfrsvrlscns n hnrapae to reappeared then and seconds, several for prey, tunnel other several place. captured the already in had down turning she continue which disappeared of all spider in however, the cases, few a lowered. place In was in turned. spinneret spinneret turned posterior spider raised leading the the while the ended, raised that wrapping was clear When spinneret posterior 3c). was (Figure trailing it the however, and lowered cases, was when some deduce with In sheet to in irregularly. the possible as against only not dabbing abdomen was by and it wrapping spinnerets initiated tunnel, tunnel. She her the the lines. of of wrapping portion mouth attached the outer made at she the prey of previous surface a irregular of the top on placed often was additional prey an new attacked she a when and behind left prey, were prey captured Previously prey. per wrapping determine to possible not was It in 3a). (Figure as sheet positions the in asymmetrical from as same whether, away the dorsally, directed in was were spinnerets posterior reduncus two spider’s the instar. fifth the approximately until common angle total (mean chelicerae her in prey the held turned she while first turned sheet), (the substrate the unl n ea owa tb unn npaewiesadn vri.Setre enof the mean inside a turned cm 1–2 She her it. or over with mouth standing while the it the place set seized at in she turning prey, sheet by seconds 788 the the it few wrap on to to a immobile, began sheet Within and nearly retreat. the tunnel, already the across was to rapidly which back run ran prey, to immediately web be then to and funnel chelicerae, as her so of water edge up the take to (perhaps feed. s Melpomene 30 to pressed about returned and for centimetres then sponge several effectively?), wet walked more a and feed before prey of to distance the piece able prey short left a the a she to of it cases mouth surfaces carried two her lateral often In and She feed. dorsal effort). to the apparent pausing on little press (with to break package. lines and single package, wrapping a the causing in placed sheet, both chelicerae, the her wrapped in then it and carried then prey, and first capture, of the 3b). site (Figure on the images it at prey video new some the in wrapped first visible were lines wrapping two least At spinneret. hl h pdrtre,setpe otnosywt e ap nadna h de of edges the near and on palps her with continuously tapped she turned, spider the While uigms rpigbhvortesie to vro eyna h rya she as prey the near very or over stood spider the behaviour wrapping most During and movements, jerky rapid her spinnerets, posterior spider’s the of length the of Because turned, she as that clear was it visibility, good with episodes wrapping different nine all In hntesie iihdwapn,sesie h ryi e hlcreadlfe tfrom it lifted and chelicerae her in prey the seized she wrapping, finished cases) spider the eight When of (seven usually spider the attacked, she when prey previous a left had she If of II stage in As ¡ 250 5 5 h edn otro pnee xeddpseiry hl h riigspinneret trailing the while posteriorly, extended spinneret posterior leading the : 982 168 u ( N ¡ ¡ p (Agelenidae). sp. 5 .reduncus P. 155 52 )drn ahbrto rpig n efre 1.5 performed and wrapping, of burst each during 6) u ,adte fe h a e h rydw nteset(enttlangle total (mean sheet the on down prey the set had she after then and ), u .reduncus P. .A h und h eidclylwrdtetpo e boe and abdomen her of tip the lowered periodically she turned, she As ). oeatcmnst h he eemd nywt h leading the with only made were sheet the to attachments some , h pdrtre npaewieatcigwapn ie to lines wrapping attaching while place in turned spider the , h pdrawy ( always spider The Psalmopoeus vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution and Drosophila N Tengella 5 0 etteetac ftetne at tunnel the of entrance the left 10) u ae nwapn dabbed wrapping in later but , p ry n i o become not did and prey, sp. ¡ 39 u , N ¡ 5 . ( 0.6 1 n1 episodes 12 in 118 N 5 )brt of bursts 4) 1641 P. Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 iue3 rpigbhvorof behaviour Wrapping 3. Figure 1642 u nafwcssas sdwapn tlatbifyt ntaeimmobilization. then initiate to and briefly later), least s at 0.07 prey, wrapping most as used for also little wrapping cases (as post-immobilization few quickly a only released was in performed prey immediately its the but thus occur): and with species not attacks did other This prey spider perhaps three wrapped. (or In the the short ant. very grasping the toward was of before bite side pulled first either legs the on ants, web and hind soldier few the turned on its to it a silk biting; within with In wrapping before it. Then, pushing briefly movements wrap wrapped chelicerae, chelicerae. to wrapping spider its began the spider several with however, the ant, it made prey, soldier the seized a on on and hold attack prey its one shifting the without to apparently and ran seconds the or with walked used spider movements the wrapping between though differences prey, perceptible different no were There taped. hibernalis IV Kukulcania legs of movements alternate with Wrapping arw)wr iil;()tesie un hl tnigoe h ry(o hw) ihtelaigspinneret spinnerets leading the dashed the and from solid with with lines shown), indicated of (not positions the prey pair between the a elapsed s. over time 0.50 (b) The standing was lowered; raised. while lines spinneret was turns trailing spider spinneret the and the leading lowered (c) the visible; while were (arrow) (arrows) raised was spinneret .Brats&W .Eberhard G. W. & Barrantes G. (Filistatidae). Pheidole eglaradiata Tengella p nswr rpe oeetniey nnal l cases all nearly In extensively. more wrapped were ants sp. n oeaeszdnmhadamtr eaewere female mature a and nymph moderate-sized One Tnelde a )and b) (a, (Tengellidae) Tortolena p c.()Tetrailing The (a) (c). sp. Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 iwo iia eune u nwihtesie’ pedsinrt r iil n t oyi itdaot90 about lifted is body its lateral and a visible d) (c, are (b); spinnerets prey spread the spider’s toward the them which pushes then in and but (a) sequence, IV similar leg a left filistatid of its with view the spinnerets the by from lines behaviour wrapping Wrapping 4. Figure aeld1 ,ad3 .3ad00 a,00 n .0s() .3s() .3ad01 d,00 e,ad00 s 0.03 and (e), s 0.07 (d), lines s 0.17 between and elapsed 0.03 (c), time s The 0.03 (b), (f). s lines 0.10 wrapping and (e) 0.03 pull IV (a), leg to s left 0.03 begins the more and toward and pull 0.03 abdomen to spinnerets (f). 3: its prepares the twists and and spider 2, near prey the the 1, brought view, from labelled posterior is withdraws a IV leg in leg this f) left (e, the as spinnerets; while prey the the from toward silk silk wrapping pushes IV leg right the ai,atraemvmnso esI;hradmnsugadicie rmsd osd in side to side a from inclined and attacking swung abdomen her When IV; very legs of protruded. not movements retreat. alternate the still rapid, were in behind often prey prey globosus Physocyclus previous antennae Wrapped the left and backward. usually were spider legs walking these the whether prey, usually their clear it subsequent not prey, the retreat, was and of prey, (it manipulation so the its compressed, further did lifted Without it toward web). simply as the it from sheet moved cases, silk the cribellate other from or In away lines mouthparts. silk wrapping of its cone to a them pulling wrapping). bringing were by IV lines legs sometimes (both cutting spider brace posterior the moved a then and of and clumsy, lack prey, for the appeared apparently of movements posteriorly, vicinity lurched wrapping not the but of to Some cases, metatarsus anteriorly some again. the inclined and in ventrally or them rearward generally moved tarsus contacted have then The was well It used. prolaterally, may abdomen others. be and sharply in would spinnerets, the that the moved near leg stroke, passed hind abdomen, leg each the this the of For side of spinnerets. the toward midline rapid the laterally the most from s; crossed The 0.03 away about legs. had only pulling lasting spider’s which spinnerets, their the the recordings; near tarsus, movement video of ventral the the positions the was in the leg visible the lines from of seldom movement Wrapping deduced were 4d). were (Figure they IV as positions legs sparse, probable spinnerets its and its of thin from strokes apparently lines slow were pulled moderately apparently alternate, and (usually) spinnerets, with its spread 4a–d), (Figure steeply fe aigwapdtepe,tesie uldi rmtewb oeie clearly sometimes web, the from it pulled spider the prey, the wrapped having After body its inclining by substrate the from abdomen its raised usually spider the wrap, To (Pholcidae). h pdrbgnatcsb pligwapn ikwith silk wrapping applying by attacks began spider The vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution uucnahibernalis Kukulcania a )I aea iw h pdrpulls spider the view, lateral In b) (a, . 1643 u ; Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 ihafl faprnl iud rsmbyguysbtne b close-up. (b) substance; gluey presumably liquid, apparently of film a with ihrsett l te pce a htlg I fe apdtepe,sometimes prey, the covered on tapped bristles’’ were ‘‘serrated often prey of were those not There the III to glue. to similar lack was legs somewhat to applied IV appeared rotation lines that tarsus lines the later first difference of but was The 5), One (Figure direction species began. glue follow. with wrapping (the other to and before difficult web, all rotated just and the extensively, to was rapid, from respect were it freed movements with was while spider’s prey applied The the consistent). were Later movements. lines these wrapping with coordination close 1644 iue5 a )Prin fa of Portions b) (a, 5. Figure rpig u ahrb ptigsik iko otepe.Sm rywr then were to able prey vicina been Philoponella have Some not would silk. we video prey. spitting as the lines, the relatively in the wrapping visible from involved sticky seldom them for to Wrapping only prey distinguish were check wrapped. on lines not then did Wrapping We IV. silk and records. legs bitten of sticky movements first alternate spitting were slow, others by while rather wrapped, but wrapping, longipes Scytodes (W. preparation). silk in wrapping Barrantes, sticky with G. associated and probably Eberhard are G. that 2004) (Agnarsson theridiids and tahdhrdaln oti ais n mre sasahwe h wn e abdomen her swung she she when when swath initiated a apparently as were emerged lines and radius, wrapping this The to rested. dragline prey her the attached which on electron web orb scanning the bifurca under Allocyclosa observed when material 2006). al. viscous without et smooth, additional were (Eberhard each but any microscope adhesive, of of movements somewhat beginning were side-to-side sign silk with the wrapping coordination any From of in Lines forthcoming). and abdomen. alternation al. the strict of et in moved Weng IV 2006; legs attack, al. et (Eberhard elsewhere .Brats&W .Eberhard G. W. & Barrantes G. (Uloboridae). (Araneidae). (Scytodidae). Drosophila l hthdbe rpe ytepholcid the by wrapped been had that fly h pdrapoce h ryaogterdu nher in radius the along prey the approached spider The h rpigbhvorof behaviour wrapping The nta moiiainwsnvracmlse by accomplished never was immobilization Initial .phalangioides P. Krhe n pebc 1990) Opderbeck and (Kirchner .vicina P. hsccu globosus Physocyclus sdsrbdi detail in described is eecovered were Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 n V h hnrahdrawr uwr ro nFgr b n ezdti etclline vertical this III seized legs and her 6b) of Figure tarsi in the arrow with (upward point rearward attachment attachment, reached the turned of then of moment and She side the IV. either 6a) At (Figure and on 6b). line (Figure leg this adhered this held prey probably the withdrew she which then to line I, sticky leg vertical one the touched The with She typical. line. times is sticky more 6 vertical 180 not Figure or long has in a one that simultaneous to depicted prey behaviour adhered fly included a which the muscid prey, once, and the a at approached gland), on IV slowly legs attack spider aggregate both The with the previously. prey described in from the of others been to balls (presumably of line visible that it wrapping had for of which reference on line application of a liquid point using a clear began as attacks serve a wrapping can Some it Theridiidae. as family detail, the in described be will species evexum it. anchored Theridion had she IV legs where the anchor with prey such to the wrapping 5–10 prey leaving Simultaneous about the feed, making to from After began line prey. moved spider the ‘‘anchor’’ IV the to an sheet, line legs attach the drag to Her one her to excursions prey attach it. lines short the to with freed alternated back from wrapping then having away of sheet, bursts lengths after Brief body prey. spinnerets or the her two near from sheet, to or silk to the carried on it that in lay movements to anterior was alternate it making in slowly, while prey relatively the either holding while prey, legs removed her the with then a ventrally spider in lasted sheet the (as bite the cases chelicerae The pushing some her prey. by In sheet the the subsided. to from struggles rapidly prey run prey’s the had the spider until the more, after or bite, minute long a with began attacks lent oeet lg edteobadssandtesie) h ieto of direction The spider). the with it sustained to on Dubiaranea and silk laid orb IV legs the it while consistent. free, held III not came and prey was I II the rotation legs After (legs from lines. and away these palps movements were ventrally cutting the by web alternate by lines or the rotated chelicerae, pushing Wrapping slowly her either often in by bitten. it was the it held to being freed she adhered as gradually by prey prey spider the the the subdued while orb, wrapping the been initiating of After had lines dragline. the prey on initiated the and sometimes Robinson after 1969; related al. only the et occurred (Robinson and species this 1971) of forthcoming). Mirick behaviour al. the et of (Weng descriptions eye previous naked the of clavipes to drops Nephila visible revealed not microscope were electron thick, that other the a from lines with under away rotated covered these in Examination soon applied) on prey was silk. were prey liquid behaviour the white the lines and of of wrapping similar rapidly, sheet the side applied opaque were (where for upper silk side of the amounts lower 1982 Large consistent: the her. and 1971, was Eberhard spider, al. rotation the et and of toward Robinson without 1971, direction 1974, spinnerets Olizarri The her her and Robinson with from araneines). Robinson radii directly (see and more lines legs or legs wrapping her Robinson one of of with on movements lines swath fashion alternate these a ‘‘rotisserie’’ with touching pulling prey prey the thus the rotated legs, to she anterior on when laid also was and Silk IV, wrapping. begin to laterally u Fgr b )t ei rpig al ntetr h pdratce e rgln to line drag her attached spider the turn the in Early wrapping. begin to c) 6b, (Figure ? p (Linyphiidae). sp. (?) Lnau,16)(Nephilidae). 1767) (Linnaeus, Nephila esrig 84(Theridiidae). 1884 Keyserling, ,o u tfe rmteset hl nsilohr h wrapped she others still in while sheet, the from free it cut or ), .maculata N. sin As vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution iyhamarginata Linyphia Rbno n oisn17) Wrapping 1973). Robinson and (Robinson l u bevtoswr optbewith compatible were observations our All h eaiu fti slow-moving this of behaviour The Kc)(brad16) all 1967), (Eberhard (Koch) 1645 Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 b,01 c,1 d,01 e,00 f,ad00 (g). s 0.03 s 0.24 and (a), (f), s 0.27 s of 0.07 was positions positions (e), segments second between s 0.12 and additional the movement first to (d), show pulled IV between s arrows legs elapsed 12 then two (small time (c), the pull She (g) The between s to lines). 0.12 IV line d). begin dotted the legs (b), and to applied (c, both solid she (c) of line Finally, the spinnerets (f). movement She by her wrapping IV ventral indicated c). to leg sticky simultaneous (b, right IV and a leg above of (e) with right line IV segment prey her leg the line brought left first the grasp her also to to the with and line line II (c), been wrapping drag leg III her have leg with attached left posteriorly may She her reached (b–e). what with and prey line the (b), drag from descending her away been seized face had to she turned by which she prey along fly Then a it. on withdrew attack quickly an and of stages Early 6. Figure 1646 h ieaanttesdsadee nt h a ufc ftepe.Lg Vte eundto returned then IV Legs prey. the of surface far the pressing to line sometimes moved on movement, even wrapping then and ventral-medial IV sides line the sticky the wrapping legs continued against of of IV two line segment legs the was The tight segment Her the 6f). movement prey. tight pressed Figure the and pulling lines a 6g), against (Figure (dotted this prey was IV When the tarsi toward there g). simultaneously extended 6g), 6f, Figure two (Figure pulled the in line and between 6f) she sticky lines (Figure prey, spinnerets of (solid the her toward length fibres near complete IV IV two additional leg tarsus least this other an moving at her III Before out included with glue. leg line line length of wrapping This one a balls the out 6d). visible with seized Figure pulled the bore body in and and of 6c), arrow f), her rest (Figure 6e, (longest the from spinnerets (Figure line for her away to wrapping herself line IV support sticky tarsus drag to of one use the brought would d), pushed she 6c, also (Figure that hold She a sequence. 6c), (Figure wrapping II leg one with .Brats&W .Eberhard G. W. & Barrantes G. hrdo evexum Theridion a h pdrtpe rywt e ih e I leg right her with prey tapped spider The (a) . Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 Vsugvnrlyadatrol oapytesik iet h ry(otdlnsi b) h ieelapsed time The (b)). in lines (b). (dotted s 0.07 prey and the (a) legs s to 0.03 both then was line and lines sticky line, dotted sticky the and held of then length apply solid additional leg by to an This indicated pulled line). anteriorly IV positions black leg and (dotted between right spinnerets ventrally the the while from swung (b)) line in IV wrapping lines sticky (solid of line segment sticky a the pulled IV leg left The h rgln stesie undpirt rpigwsntceri h ies Sticky videos. the lines in non-sticky tesselata clear before Achaearanea s) not 43 to was (up wrapping time to long unusually prior applied. an turned were for applied spider were the lines as wrapping line drag the of stages later the In prey. wrapping rufipes IV Nesticodes the leg simultaneous. most to than and rather sticky, line not alternation probably wrapping in were were that of movements lines segments applied spider further the wrapping, apply and seize by behaviour attack early IV legs of use simultaneous in Stages 7. Figure enprilyrsrie,tesie rdal u h erylnso h he hl she while sheet the of had lines prey nearby the was the After line cut wrapping IV. gradually sticky legs spider the of species the movements both restrained, alternate lines In partially these and sheet. eye); been simultaneous the naked on both the by place using with attacks in noted applied initiated prey (easily species rapidly the both liquid ran Nevertheless, held with spider it. laden the generally her attack while heavily webs to and sheet, lines sheet spider the their with the of the wrapping of in surface between surface upper sheet barrier lower the of the a on horizontal those on formed generally dense from was sheet prey different the a The and quite prey. construct forthcoming), was species al. occurred et both species (Eberhard two because these theridiids, by other attacks which in context with line wrapping sticky applied and in away, as face IV to legs her turned of spider movements simultaneous the repeated legs, front her with Lcs 86 (Theridiidae). 1846) (Lucas, Kyeln,18)and 1884) (Keyserling, vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution Chrosiothes fe prahn h ryadtuhn it touching and prey the approaching After .evexum T. p nr. sp. etcie ruficeps Nesticoides Fgr ) h oiinof position The 7). (Figure porteri (Theridiidae). namsodfy (a) fly. muscoid a on 1647 The Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 nlsmsstudiosus more movements Anelosimus IV leg lines later alternate wrapping the utilize multiple In to thinner, tended prey. use also the to and to frequently. liquid, appeared access obvious species any greater both lacked lines of that spiders wrapping attacks, her of giving stages thus wrap, to continued eldu hsln gi nhrwybc ohrrsigpae ntertr rp she trip, return the on place; 180 turned resting she her place resting to her reached back she way When line. and her sticky broke new then on with and again these line prey, the the with line approached replaced accord she this broke which in spider up along were the line that reeled spider horizontal in the theridiids second up other a reeled of that and of simultaneous from differed videotaped observations with behaviour one applied Direct Attack In was observations. IV. line. and this legs sticky, of apparently of end was one movements line at wrapping rested first spider her the sequence, and line, horizontal IV. sticky legs single of spider movements reimoseri the alternate attack, consistently Phoroncidia used an stages often in early and Later silk the IV. non-sticky legs and of appying biting, movements to than simultaneous shifted rather and wrapping silk sticky with included began usually flies on great with ants pacificus these A. treat on to attacks appeared spiders in The IV. only was caution. legs silk of biting wrapping Sticky movements IV. to simultaneous legs of prior movements alternating using used bite, the following silk dry studiosus tcysl n moiiainwrapping, immobilization and silk sticky algtilmnto.Tesie prahdtepe lwy apn ha ihher 180 with approximately ahead turned using tapping she under long prey, filmed slowly, others the the five prey were touching and of the After attacks lines), most I. approached silk built Two legs the spider covered spiders long forthcoming). of The that The images al. illumination. material occasional species. daylight et provided sticky this 2004) (which (Eberhard of of light al. infrared lines behaviour amounts et the capture large Arnedo of vertical relatively 2004; description with (Agnarsson detailed webs Theridiidae more these a and of justify Nesticidae All between IV. legs relations her of calidus movements Gaucelmus alternate of with behaviour the her to behind identical line were the details in reeled and 1648 o aebe oee ihvsi aeil h rpigmvmnso h esI during IV legs more the IV of movements may legs wrapping they The of that material. suggesting movements viscid videos, with the alternate covered in using been visible have consistently gradually not less but were lines IV, These legs frequently. the moderately attack with with an line applications covered in after This were soon lines silk. spider lines 8b). Figure the viscid many (dotted from microscope; in confirmed of taken a IV was balls lines were under suggestion small that legs checked (dotted This prey and her bundle three balls. initiated on sticky was prey of lines pressed carried the the was each probably examining line of it by of the sides that that stroke so suggesting or simultaneously gleamed, a bottom legs these with the moved against rapidly spinnerets then her and 8a), Figure from line silk h pdrte ple diinlwapn ik tfrtmsl sn simultaneous using mostly first at silk, wrapping additional applied then spider The takdmn ucdaddoohldfisb iig n nywapdte with them wrapped only and biting, by flies drosophilid and muscid many attacked .Brats&W .Eberhard G. W. & Barrantes G. ei 96(Theridiidae). 1956 Levi, etc,17 (Nesticidae). 1971 Gertsch, ei 94(Theridiidae). 1964 Levi, Hnz 80 (Theridiidae). 1850) (Hentz, ncnrs oterlcac of reluctance the to contrast In .pacificus A. .studo P. Pseudomyrmex h netit eadn h phylogenetic the regarding uncertainty The h e fti pce osse fa of consisted species this of web The Eehr 1981). (Eberhard ossetyue oh hi attacks Their both. used consistently auefmlso h solitary the of females Mature ns n a ple with applied was and ants, u n uldasticky a pulled and , A . studiosus outilize to A u . , Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 esI n ntae h takb unn prxmtl 180 approximately turning by attack the initiated and this of I, portion legs distal the silk. case, viscid one Synotaxus with least covered at apparently In and thread. shiny capture unusually new began. was the a feeding line been after before have often web may the wrapping, which carried of of was tangle s prey 45 upper The to returned. the then even in up and the sometimes retreat after retreat of IV her the side until toward tarsus to far bitten upward and the away not and spider, moved was sides had the spider the prey from around The prey lines package. the wrapping of prey apparently thus side midline, far its the crossed of to midline silk the the spinnerets. across applied barely the only near moved line tarsus the the of grasped tip and the spider that the in unusual were stage this between line was sticky lines the dotted and snagging solid prey, by the indicated past positions IV between elapsed legs female time drawings. both The both moved mature (b)). in then in a s and lines 0.1 of (dotted (a)), prey of IV the lines on legs dotted them of in line movements pulls wrapping IV simultaneous of view calidus Antero-lateral 8. Figure hr h idtru otce t n hswsntgipdb igesrcueo the on structure single a point by the gripped at not widened was often thus tarsi swath and two The the it, common. between contacted and and leg. tension tarsus clear them, under hind were on lines the 9), wrapping glue of (Figure where Simultaneous have swath prey droplets. to a the scattered applied seem to small, which not IV very IV, did only legs lines had of The chamber movements white. humid a in in a prey lines only the coated had quickly each that two other them. affinis the on Azilia humid and liquid a balls, of in a sticky being balls lacked after afford have ball. microscope entirely to tiny to a one appear single enough under this: not slow packages did confirmed prey were which box wrapped movements lines, three The wrapping of the Examination IV. of legs view both good with simultaneously prey utbfr edn,tesie oeie ecne otesbtaeadatce line a attached and substrate the to descended sometimes spider the feeding, before Just IV legs of movements wrapping Many wrapped. being while turned not was prey The h pdrfrtple tcysl wt ml lc osi rwn)fo e pedsinrt scn leg (second spinnerets spread her from drawing) in dots black small (with silk sticky pulled first spider The . p (Synotaxidae). sp. (Araneidae). pdr takdb rpigrpdywt hc wt flines of swath thick a with rapidly wrapping by attacked Spiders h pdrapoce h rysol,tpigwt her with tapping slowly, prey the approached spider The vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution u n pligsl othe to silk applying and Gaucelmus 1649 Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 h rpigmvmnsof movements wrapping The functions and details Behavioural (b), s 0.10 (a), the Discussion s 0.10 from was silk lines them pulled dotted between and IV line solid leg the by (b) press indicated (c). to other positions s (c)) between the 0.03 in elapsed then and (arrows time and ventrally The simultaneously (a) prey. female moved the one mature were against First legs a both orb). of then horizontal and movements the spinnerets, wrapping above simultaneous from of (view (stippled) view Ventral 9. Figure 1650 tec rpigsl costepe.Aohrdfeec a h eua zig-zag regular the was difference Another prey. to body the her of her across by movements produced turning silk silk the wrapping on lax chelicerae. wrapping of relying her mass from than the prey stretch rather into the prey spinneret, release the not directed moved did dorsally spider thus this the she wrapping, turned, during of she II spinnerets As stage their of part of first positions the the (Nitzsche but pisuarids prey substrate, and the the unknown. 1974) when to are Knost attacks prey behaviour and feeding subsequent fasten (Rovner or it facilitate also transport Lycosids as also 1988) facilitates chelicerae). (especially may perhaps the it thus wrapping in compact and theraphosid, remained to site), the or Wrapping wrapping silk. substrate in the of the in (and from swath able to turned lax were away prey more we spider lifted the a where was the anchor produced theraphosid, to while the spinneret in raised functions substrate least the apparently the at details, tightly assumed and more dabbed less raised, observe spinnerets repeatedly or was to striking it more spinneret lateral most a other as posterior The The produced silk place. observed. and the of lowered we that lines was swath species PL was pulling stretched other One by species the positions. than in asymmetrical three rather similar, as these away, to IV abdomen pulling among legs the then similarity with and and pulled spinnerets spinnerets substrate was the the Silk the dabbing from immobilized. to by and attachment spinnerets bitten an (PL) being make after lateral substrate posterior the the to from fastened was prey the h hrpoi ifrdfo h te w pce ntobscrset.Frt during First, respects. basic two in species two other the from differed theraphosid The .Brats&W .Eberhard G. W. & Barrantes G. .reduncus P. , .radiata T. and , Melpomene zlaaffinis Azilia p eesmlri that in similar were sp. rpigaprey a wrapping Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 htalo eryalsiescntutasl oeigfreg Fei 96,adseveral to and functioned 1996), originally spiders) (Foelix for eggs synapomorphy for (a covering lines silk silk a that construct appears argued spiders It have all construction. egg-sac authors nearly is or behaviour all wrapping of that origin evolutionary wrapping possible prey One are of origin alternatives the some and construction and Egg-sac hypotheses The transitions. evolution, of parsimonious their most numbers the in being below. the not roles discussed of important to sense the play respect in probably ‘‘orthodox’’ Because with not obviously traits conclusion. are figure definitive hypotheses of our a this values of families than and some selective rather spider data, that work of among believe future lack for we relations for guide omitted speculative phylogenetic a are the represents families 10, concerning Most Figure 2005). in hypothesis illustrated (Coddington recent are observations a our by using suggested hypotheses evolutionary Several hypotheses Evolutionary absent. rmaiiomgad)wr en ple otepe.I diin hyocre nthe in occurred they addition, In prey. (presumably the glue to of applied balls in obvious being without synotaxid were early glue. lines glands) with common thinner covered aciniform when especially lines later, from wrapping were occurred applying also IV were they nesticid legs But the the with and that theridiids applications important. when suggests Simultaneous probably wrapping caution is strict. This distance not prey. safe was the a of at range while attacks of wrapping out beginning slightly of still advantage was spider the when Gaucelmus seilytgtrsritt vroepe tuge Eehr ta.2006). al. require the et not (Eberhard does and sticky struggles function bundling prey restraining the simultaneous the overcome when the contrast, from to attack, In restraint an Both occur. tarsus tight in to difficult. early especially tended the important IV particularly be legs of to of be likely movements Removal more may the are preparation). theridiids of functions short extraction in of fell G. line Barrantes, silk (W. lines wrapping wrapping wrapping G. the from of when tarsi alternating and Facilitation their occur removing another. IV even by Eberhard problems one could frequent legs against have line pull prey spiders both the could Some from tarsi the it prey. with two after leg the silk the toward lines as wrapping of of prey, wrapping pushed removal segment the new to of each sometimes applied from been application tarsi has are the Simultaneous extricating Simultaneous that facilitates IV. also they wrapping. probably lines compared legs when restraint of lax tighter tight of effective, were more movements more stages lines provided have wrapping the initial may The and with related prey, functions. the the closely possible to the during two applied and were least theridiids species at in apply has synotaxid however, to application movements occurred, IV and reliably leg prey Simultaneous nesticid the alternation. to strict silk in moved wrapping usually more IV produce species legs glands prey, two piriform other their The because 2005). 2a). perhaps Coddington (Figure (see movement, began attachments of secure wrapping type this when made substrate never the to attachments h soito ewe iutnoswapn oeet n tcywapn lines wrapping sticky and movements wrapping simultaneous between association The eea pce,icuigtetheridiids the including species, Several wrap to spinnerets their from silk draw to IV legs their used which species, other the In Synotaxus n h uloborid the and , p n h araneid the and sp. Philoponella vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution zlaaffinis Azilia hwdcuinb ntaigwapn attacks wrapping initiating by caution showed , Theridium nwihsik ie eesas or sparse were lines sticky which in , and Nesticoides h nesticid the , 1651 Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 esrig 83(.Eehr,upbihd.Seisi te russc sthe as such groups other ( in Species Mello-Leita unpublished). uloborid Eberhard, the clubionid (W. and unpublished), 1883 Barrantes, Keyserling, (G. 1894) (McCook, bevddrcl nti td ri rvospbiain * natnaiepyoeei re(oiidfrom (modified groups tree the phylogenetic of tentative behaviour a text. the the on of in (*) super-position discussed publications are web by previous immobilization hypotheses the illustrated the on in versus are of it or Justifications rotating hypotheses wrapping study 2005). including by The Coddington this non-immobilization prey snared. characters, in the IV, was wrapping directly behavioural and legs it observed IV, prey-wrapping which moving legs on of several versus movements line of simultaneous body versus evolution the alternate wrapping, moving the by regarding wrapping Hypotheses 10. Figure 1652 aiuaelnswt hi esI nohrcnet.Teeicuetearaneids the spiders include which These in contexts. groups other in in IV occur citricola legs and substrate, their the species, with with and several lines contact in into manipulate drawing brought construction Both probably spreading egg-sac are and above. during spigots abdomen, the occur describe the that and we so spinnerets apart that the spinnerets of the IV movements legs similar dabbing in are using using trait species lines some ancient applying without of extremely behaviour construction an egg-sac wrapping the probably in to used is patterns construction behaviour The Egg-sac spiders. 1964). (Kaston eggs cover atpoadizzydeani Mastophora lobidegsc ydbigteadmnt h usrt.O h ai ftewidespread the of basis the On substrate. the to abdomen the dabbing by sacs egg build also 5 Trachelea (Forska .Brats&W .Eberhard G. W. & Barrantes G. goc brunnea Agroeca o 94(ice n acnelsNt 05,adtepisaurid the and 2005), Vasconcellos-Neto and (Fischer 1934 õ, ) ornata l 75 Klmn 1961), (Kullmann 1775) ˚l, (Mello-Leita brad 91(brad18b,and 1980b), (Eberhard 1981 Eberhard, Bakal 83 Hl 90,tesicariid the 1940), (Holm 1833) (Blackwall, o 93 L .CsaShit esnlcommunication) personal Schmidt, Costa E. (L. 1943) õ, rnu quadratus Araneus Gtz 89 Coe1956), (Crome 1889) (Getaz, Metazygia lbrstrilineatus Uloborus ooclsintermedia Loxosceles prob. Paratrachelea Cyrtophora wittfeldae (?) Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 r abdt h usrt,w rps httewapn oeet of they movements when wrapping spinnerets the the that from emerge propose that we silk substrate, of the made to sacs dabbed egg are of spiders in occurrence radiata otepe a vle ovretyi e-uligseisi tlatfu n perhaps and four least at in silk species wrapping web-building apply to in IV convergently legs evolved of has use prey the the then homologous, to is Agelenidae IV. and legs Tengellidae, with prey silk fact the wrapping the over Pull to and past due lines. run part to drag in needed in are is it lines convergences tangle manipulating these and in Perhaps dexterity prey. additional little the other that over several and in around independently, rapidly (Dippenaar-Schoeman probably Hersiliidae 1967), Jocque evolved, (Glatz also and Oecobiidae has including IV groups, legs entelegyne of involvement direct of movements and positions no and as however, attachments certain, of not sequences spinnerets. is the This regarding 1990). given Ketner were and details (Coyle to legs retreat’’) the of and/or diplurid to use surface returning the without prey after in behaviour the occur Wrapping to 140). also p may spinnerets 1979, IV the Lubin and from (Robinson directly substrate’’ silk the attaching insect, the for around here reported three behaviour the our the from to for similar prey problem of a loss not prevent was to (which adaptation 1974) an Knost lifting be and species). when may (Rovner lines is This sites wrapping here wrapped. wrapping break described was not elevated behaviour did it positions, spiders after the spinneret these prey with of that the behaviour was details this difference note and palps One not of the wrapping, certain. did homology process not of Knost possible the and part Throughout the Rovner first turn. so prey. to the however, the continues during contact it to chelicerae while the used substrate to its are the silk with on attaching prey it repeatedly leaves the turns, then and holds prey also the It over substrate. stands arose spider groups The two respects. so the several between seem similarity distributions this such that convergence. suppose of adaptive to to were consequences credulity spinnerets due asymmetry selective strains the if Such it likely expected that spinnerets. that the small from PL but package is of prey symmetrically; the use silk, is in on used asymmetrical derivations, manipulate lines wrapping of their independent to distribution in to non-immobilization the legs alters similarity opposed of hind the as homology by spiders, the supported agelenid The using and without spiders. tengellid, spinnerets, mygalomorph, araneomorph moving the by in wrapping and prey ancestral body that its propose entire and we families, behaviour, the spider construction other egg-sac several occurrence to of its resemblance behaviour spiders, wrapping mygalomorph post-immobilization in the occurrence during its body. capabilities), behavioural the of of number movements using silk silk wrapping glands. wrapping Pull known epiandrous all the because than likely rather less by spinnerets produced seems the is spigots, from that their web comes sperm and the glands silk, epiandrous of source the ancient similarly altenative, An behaviour. h s fbd oeet natc rpigt uladapysl otepe without prey the to silk apply and pull to wrapping attack in movements body of use The psechrid The of species two of behaviour wrapping The and , 97,adGahsde(rsoe15) nteegop h pdrruns spider the groups these In 1958). (Bristowe Gnaphosidae and 1997), ´ Melpomene Fenecia p rp paetyimblzdpe nawyta a equite be may that way a in prey immobilized apparently wraps sp. p ecie eewr eie rmegscconstruction sac egg from derived were here described sp. ftewapn eaiu nteseiso Theraphosidae, of species the in behaviour wrapping the If Ischnothele vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution eglaradiata Tengella ‘apyn ikt h era alimmediately wall retreat the to silk (‘‘applying Lycosa ntebsso t ipiiy(ntrso the of terms (in simplicity its of basis the On Rve n ns 94 ssmlrin similar is 1974) Knost and (Rovner h pdr‘bnspe ycircling by prey ‘‘binds spider the : .reduncus P. 1653 , T. Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 1654 rpigw ecie bv,i hc n e Vsie ikbtte assfra for pauses with then simultaneously but it applying silk before seizes it ‘‘simultaneous’’ IV seized 6–8). also (Figures leg the has prey one IV the from to leg which both second differs in the until above, description moment described This we movements’’. wrapping wrapping these attack synchronous of with stages theridiids early nearly linking the in synapomorphy araneid that a noted the (1971) be by the Olizarri in may and occurrence study Robinson its families. and this simultaneous two derived, in apparently comparisons, is synotaxid outgroup lines of and wrapping basis nesticid apply the to On IV legs known. simultaneous of not versus movement are alternate IV regarding also legs details of and but movements 1978), observed, (Carico we silk adhesive-laden theridiids with of species 12 the and of 1980a) theridiids, all (Eberhard legs other both in two with occurred in silk silk wrapping attacks of occurs of sheet in application horizontal Simultaneous early dense spider). a IV the to and lines prey non-sticky the or Garcı between sticky also single (from (see occurred are mostly wrapping IV applied and silk legs non-sticky of phylogeny, of prey. evexum use to movements later this lines IV, alternate wrapping legs in apply with of to movements hypothesized IV simultaneous with legs is mostly its used than that and ancestor related pholcids common that a closely is from alternative drymusid more descended One The 1974). are 1967). (Valerio prey (Eberhard prey its scytodids web the rotating the against by from repeatedly spinnerets freed spinnerets its is from prey silk the dinora pulls after Drymusa instead legs and anterior IV, its legs with its with directly lines omncto) nte theridiid, Another communication). elkonvraiiyo takbhvor(oisnadOiar 91 ui 1980; Lubin 1971; Olizarri in and spiders diguetid (Robinson The was that wrapping behaviour fact that attack fact the Japyassu the involve of from by not comes variability do also IV) well-known Support that ochyroceratid legs IV. web-building wrapping legs the with of with in absent prey styles lines not the different does release to that utilize silk and abdomen applying each the seize of Dyrmusidae use and to direct if Diguetidae more and ability to correct, ‘‘regress’’ and is the not Pholcidae 2005 does in require Coddington IV wrap by legs to presented alternate IV phylogeny of legs use the of (if use supported the is of Scytodidae derivation and Independent 1998). these the orientation Huber so 1990; lines, from complex silk release silk involves then sticky and struggling) traits. apply, by pull, still multiple grasp, involve immobilized is to convergences immobilization IV been prey legs (especially of had wrapping the movements coordinated effective it while Pholcidae (1975), initiated after initiated Robinson after); was but wrapping by (wrapping others bitten Scytodidae noted in and As was before); prey); biting, chelicerae). prey slightly the before occasionally biting the to initiated but before prior bitten was initiated was Filistatidae was wrapping prey 1969); (wrapping some Shear the prey by and (in after the observed after initiated 1983) Orbicularia was initiated was wrapping is (Gray no wrapping wrapping Hypochilidae Gradungulidae (most families (in both bitten) include (in been 2003) These has al. et lineages. (Lopardo araneomorph Austrochilidae different five h eaieuiomt fteiidatc eaiu—al s fsik ikapplied silk sticky of use behaviour—early attack theridiid of uniformity relative The s flg Vt rpi rbbywdsra npocd Krhe n Opderbeck and (Kirchner pholcids in widespread probably is wrap to IV legs of Use n arsfrhoig evsoe h osblt hti a oeie occur. sometimes may it that possibility the open leaves forthcoming) Caires and ´ nBai)i tiigi iwo h ait fpyia otxsi hc attack which in contexts physical of variety the of view in striking Brazil)—is in .Brats&W .Eberhard G. W. & Barrantes G. iutaalbolineata Diguetia rip argentata Argiope aei,17 loplswapn ikwtotuigaylg,b abn its dabbing by legs, any using without silk wrapping pulls also 1971 Valerio, caaae digitus Achaearanea Ariamnes Fbiis 75 ‘ohlg Vpc psl n ati in it cast and silk up pick IV legs ‘‘both 1775) (Fabricius, O .Cmrde 85 osntmnplt wrapping manipulate not does 1895) P.-Cambridge, (O. uypsfunebris Euryopis Ochyrocera ( 5 ukpadMrus(.Japyassu (H. Marques and Buckup Argyrodes huhorsalsml ieadthe and size sample small our though , Hnz 80,as rp ry(ants) prey wraps also 1850), (Hentz, aadJapyassu and á ) attenuatus .P-abig 1881 P.-Cambridge O. 05on 2005 ´ ,personal ´, Theridion Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 rnissc as such araneids h yohssta es oiotlsetwt h pdrbnahi n h ryabove of behaviour prey the the by and supported it 1980). is beneath wrapping Lubin spider immobilization 1975; the of with (Robinson loss sheet favour which struggles may horizontal sheet, prey’s it dense the the a of because from that risky hypothesis spider less The wrapping the be by protect would afforded attacks protection wrapping largely biting in apply would a increase because to the reduced, such attempts addition, be to with In also barrier 1980). wrapping may a dense (Lubin forms Immobilization prey a sheet sheet. the of the to the because underside lines effective the above less on is be moves would prey spider web the the while prey: sheet, their from horizontal them isolate partially in which reduction secondary a of hypothesis the supports strongly Avile (L. n fsvrlseisof species several of and but clear; not congeneric are the species by this wrapping in immobilization wrapping of immobilization use of consistent loss the partial the for reasons The TbeI;Lbn18) lhuhte owa oeptnilydneospe (Lubin prey dangerous wrapping potentially than in some rather attacks wrap biting wrap by do see to sheet in they failure the than although our under rather 1980; 1980), from sheets Lubin prey horizontal II; attack dense (Table generally build they but and 1971), orbs, Olizarri and (Robinson wrapping es oiotlset opromimblzto rpigo ouaigmls which males, copulating on wrapping immobilization perform of do females sheet, adult that horizontal is dense idea a this fits that detail another ryecps(e oisn17;Lbn18) moiiainwapn a also has wrapping Immobilization 1980). increased also Lubin group, and another forthcoming), still 1975; al. in reduced et Robinson strongly (Weng become (see recovered secondarily not escapes and expended such prey costs, is has that also silk probably wrapping as Nevertheless, araneids. gasteracanthine and spiders, ofre yatcigpe ywapn hmi ik(oisne l 99 oisnand Robinson araneid 1969; specialized al. the et (Robinson tetragnathids, silk linyphiids, in in them 1971) wrapping Mirick by prey attacking by conferred needed not is mystery. is work this agreement Further of up observations?). lack clear imprecise this to prey? for different reason in differences? The of (species material. species clear Barrantes, gland) two (aggregate (probably) or sticky G. one with of covered observations and different in five in Eberhard also Synotaxus failed we and G. but 722); 2004) p groups. W. 2003, (Agnarsson other of theridiids to 1990; those extend in although not Opderbeck evident may is also and preparation) that (Kirchner IV the tarsus pholcids in of surface lines, ventral wrapping and paragraph), sticky previous to (see nesticids prior and in theridiids apparently and independently once, synotaxids of evolved ancestor common have may be to movements likely A that seems behaviour show (F. wrapping data weavers this so molecular theridiids, Preliminary to related convergent. closely not is genus This soitddrn h nta tgso h tak fteiid,btte r apparently the are in they but occurred theridiids, movements of wrapping attacks IV the leg araneid, of Simultaneous stages independent. initial evolutionarily the during associated h oso moiiainwapn nlnpid a eascae ihterwebs, their with associated be may linyphiids in wrapping immobilization of loss The h rpsli iue1 eursterpae oso h rbbeadvantages probable the of loss repeated the requires 10 Figure in proposal The the on setae serrated of comb the and lines wrapping sticky between association The iutnosmvmnso esI n h s fsik rpiglnswr often were lines wrapping sticky of use the and IV legs of movements Simultaneous s esnlcmuiain,a ela yseisi te hrdi genera, theridiid other in species by as well as communication), personal ´s, zlaaffinis Azilia ocnimtecam(rsode l 98 htsntxdwapn ikis silk wrapping synotaxid that 1998) al. et (Griswold claim the confirm to ´ vrzPdla esnlcmuiain.I u,smlaeu wrapping simultaneous sum, In communication). personal lvarez-Padilla, Argiope Azilia Synotaxus Fgr ) pce hc i o rdc tcywapn lines. wrapping sticky produce not did which species a 9), (Figure gi ihu tcywapn ie Fgr 10). (Figure lines wrapping sticky without again , htbidob Giwl ta.19)aduiieimmobilization utilize and 1998) al. et (Griswold orbs build that Cyrtophora ‘hr itesmlrt ihteteiidcm’ (Agnarsson comb’’ theridiid the with similarity little ‘‘share vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution Lbn18) hs pce r lsl eae to related closely are species These 1980). (Lubin Kapogea a edet u ml apesz) Still size). sample small our to due be may Synotaxus Azilia ytpoacitricola Cyrtophora sdfiutt lc mn orb among place to difficult is nlsmsstudiosus Anelosimus .studiosus A. p a h ertdbristles, serrated the has sp. .pacificus A. Chorizopes hc lomake also which , . and htpeson preys that Kapogea TbeII). (Table .eximius A. 1655 sp. Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 rm .15.Kkna n ibaeengrKezpneatndsGenus des Kreuzspinnenarten einiger Eiablage und Kokonbau 1956. W. Crome ugsin,adSR n h nvria eCsaRc o iaca support. financial for References Rica Costa de Universidad the of work, and evidence field STRI arranging and for and help Hormiga checking suggestions, invaluable G. for for Japyassu Machado Opell, Weng Glauco Brent Julin Hilton lines, wrapping Levi, specimens, nesticid W. identifying on H. kindly glue of Huber, for Bernhard Ubick loss Agnarsson, Darrell Ingi subsequent thank We however, require, Acknowledgements would hypothesis this a on in derivation Even based wrapping independent were immobilization an with conclusions araneines. accord in their acquired is (apparently This in wrapping). that was deinopoids related rotisserie only (1997) of Coddington and in definition and and and tetragnathids different Scharff present of in theridiids is Orbiculariae, claim the occurs wrapping to 1971), of it contrary Rotisserie (araneids), tetragnathoids. Robinson groups ancestor these some of and one and common araneids, (Robinson some the deinopoids families, in in immobilization that absent orb-weaving independently suppose to number the was be the reduce would in would wrapping wrapping that immobilization ways. 10 wrapping of Figure these losses in in evolutionary immobilization scheme explained of the to be of alternative cannot parsimonious loss 10) less (Figure attack One the araneids surprise gasteracanthine its But and and venom, tetragnathids 1983). strong apparently (Eberhard chelicerae, large tactic its to related presumably rpiglnso iyhist eal orsri h rydsietebriro h sheet. the non-sticky of the barrier the are despite than prey likely the more restrain to thus able are be and to linyphiids glue, of abundant lines with wrapping coated were lines paykulliana ol A enrN.19.Osrain ntepe n rycpuebhvoro h unle mygalomorph funnelweb the V, of behaviour Roth capture prey PE, and prey Cushing the on P, Observations 1990. Paquin ND. Ketner D, FA, Ubick Coyle In: spiders. of classification and Phylogeny 2005. JA. Coddington in behavior Norton. Predatory London: 1978. spiders. JE. of world Carico The 1958. WS. Bristowe O zur Untersuchungen 1972. relationships Araneoidea, R. phylogenetic Blanke tree: (Araneae, a to relatives comb a their From 2004. and RG. Gillespie spiders I, Agnarsson cobweb JA, Coddington of MA, Arnedo phylogeny Morphological 2004. I. Agnarsson sheet horizontal dense under immobilization from Effective 1972). theridiids (Blanke three female by the as performed webs, sheet were the attacks of side wrapping same the on occur 1656 gaso .20.Tepyoeei lcmn n icmcito ftegenus the of circumscription and placement phylogenetic The 2003. I. Agnarsson h oso moiiainwapn noeaaed h pdrpredator spider the araneid, one in wrapping immobilization of loss The noooiceZishit(F 3:28–55. (NF) Zeitschrift Entomologische dtr.Sieso ot mrc:a dniiainmna.Aeia rcnlgclScey www. Society. 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(Araneae, , .studiosus A. Chrosiothes ¨ ohsooi n O und kophysiologie uypsfunebris Euryopis . p nr. sp. Hnz Aaee hrdia)adteevolutionary the and Theridiidae) (Araneae: (Hentz) ¨ porteri ehlgevon kethologie n also and , ytpoacitricola Cyrtophora Steatoda Synotaxus ( Chorizopes 5 Araneus Forska Lithyphantes Deutsche . l(Araneae, ˚l (Araneae: p,is sp., ) Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 ryM 93 h aeof male The 1983. M. Gray lt .16.ZrBooi n opooi von Morphologie und Biologie Zur 1967. L. Glatz olxR 96 ilg fsies e ok xodUiest Press. University Oxford York: New spiders. of Garcı Biology 1996. R. Foelix of fecundity affecting Parameters 2005. J. Vasconcellos-Neto ML, Fischer eiH.19.TeAeia r evr ftegenera the of weavers orb American The 1997. HW. Levi eiH.18.Teobwae genus orb-weaver The 1980. HW. Levi ulanE 94 eeEgbis u Ergebnisse Neue 1964. E. von Kullmann Eierkokonbau Der 1961. E. 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WG. Eberhard bradW.18b h aua itr n eairo h oa spider bolas the of behavior and history natural The 1980b. WG. Eberhard bradW.18.Bhvoa hrcesfrtehge lsiiaino r-evn pdr.Evolution spiders. orb-weaving spider, assassin of an classification of behaviour higher Predatory the 1983. WG. for Eberhard characters Behavioral 1982. WG. of web Eberhard line single The 1981. WG. Eberhard ipna-comnA,Jocque AS, Dippenaar-Schoeman ui D 90 h rdtr eairo ytpoa(rna,Aaede.Junlo Arachnology of Journal Araneidae). (Araneae, Cyrtophora bei Beute der of Umspinnen und ‘‘Brautgeschenk’’ behavior 1988. ROM. predatory Nitzsche The 1980. YD. Lubin aC Japyassu C, á opooi e ir 61:185–214. Tiere der Morphologie esrig 84(rna,Teiide.BoaNorpc 51A. Neotropica Biota Theridiidae). (Araneae, 1884 Keyserling, Aaee iaide.Junlo rcnlg 33:670–680. Arachnology of Journal Sicariidae). (Araneae, 93:251–254. Naturwissenschaften wrapping. by h uemo oprtv olg 149:1–74. Zoology Comparative of Museum the Glenognatha 2:41–122. 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(Araneae: Mexico of North Tetragnathinae subfamily the of .19.Arcnsies nietfcto aul rtra ln Protection Plant Pretoria: manual. identification an spiders: African 1997. R. ´ rgaugl carraiensis Progradungula brdnNtbuuddsSxavratnengrSpinnenarten einiger Sexualverhalten das und Netzbau den ¨ber e hti o sae sce86:407–413. Psyche asnare. not is that web a : vlto fpe-rpigbhvori spiders in behaviour prey-wrapping of Evolution Mecynogea hrnii studo Phoroncidia ytpoacitricola Cyrtophora eoisannulipes Oecobius h ufml eia n h genera the and Metinae subfamily the , eyoe,Manogea Mecynogea, ose n ry(rna,Gauglde with Gradungulidae) (Araneae, Gray and Forster hlpnlavicina Philoponella ei(rna:Teiide:ape attractant? prey a Theridiidae): (Araneae: Levi ctdsintricata Scytodes Chorizopes Forska ua Aaee eoide.Zishitfu Zeitschrift Oecobiidae). (Araneae, Lucas iar mirabilis Pisaura ooclsintermedia Loxosceles l(rna,Aaede Zoologische Araneidae. (Araneae, ˚l , p Aaede,adtedefensive the and (Araneidae), sp. Kapogea pdr ageadkl hi prey their kill and mangle spiders Aaee ctdde.Revue Scytodidae). (Araneae, atpoadizzydeani Mastophora , and ooee fimbriatus Dolomedes Cyrtophora Mello-Leita hrdo evexum Theridion .Zishitfu Zeitschrift ). Pachygnatha, (Araneae: o1934 õ Pholcus 1657 p n. sp. und , ¨r ¨r Downloaded By: [University of Pittsburgh] At: 05:05 30 September 2007 eesH.13.WieeUtruhne u Untersuchungen Weitere 1933. HM. Peters eesH.13.DeFnhnln e rusin ( Kreuzspinne der Fanghandlung Die 1931. HM. Peters 1658 oisnM.16.Peaoybhvo of behavior Predatory 1969. MH. Robinson oisnM,LbnY.17.Seilssadgnrlss h clg n eairo oeweb-building C, some Beer of behavior G, and Baerends ecology In: the spiders. generalists: and araneid Specialists in 1979. behaviour YD. Lubin predatory MH, of Robinson evolution The 1975. MH. Robinson onrJ,KotS.17.Ps-moiiainwapn fpe ylcsdsieso h ebcosstratum. herbaceous the of spiders lycosid by prey of wrapping Post-immobilization 1974. SJ. 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