HARVESTMEN HARVESTMEN The Biology of Opiliones Editedby Ricardo Pinto-da-Rocha Glauco Machado and GonzaloGiribet llurvurdUmvcrsuvPrl'SS ClIlI/IJrill!Jl.',\1/1SSIIc/IlISl'lIs,I/lld/.0//1/11/1.FII!/11/1111 .!O1l7 \"'I",,~IIl' '::'MI~I'\Ilh·l·.·....i..klll.IIl.lldl,,\\,.,r11"n.mlt·"Il".".'· \lIn;:III,...·,...nnl I'ruu....1'11,1,..1"i"·,I"'.'k,,,1,\'m..-i.-;, I'. ,'m hh·llI.t.....I"I'ri,~,:r;,l'hi.·;,lr..r-.·r...n-.....;lI"limk, l"n'·1i:·)~:-.ll-',;--HI.,!j-Ji-;-,;,II...I';'I,,·n ISI:'·I'I:II·";-·I·'I':::I:··I';I'I.·I\;'I""-' I.(ll'ili"n,", 1.I'illl"·,I;,·I\"dlil.lU'·;lI·,I,, II \1;"·11,"1,,.1:1;1["'''. Ill.t:jril,,·l. Contents Contnbutors vii Preface ix 1 WhatAreHarvestmen? I GlaucoMachado,RicardoPinto-da-Rocha,andconzaioGiribet 2 ~lorpholograndl'uncttonalAnatomy I.J JeffreyW.ShultzandRicardoPinto-da-Rocha 3 PhylogenyandBiogeography 6] conzoioGiribetandAdrianoB.KUry -l Taxonomy ss RicardoPinto-da·RochaandooozaioGiribet 5 Paleontology 2-11 JasonA.Dunlop (, Cytogenetics ]66 NobuoTsurusaki 7 Erology .!."iu DavidJ.CurtisandGlaucoMachado VI contents I; DietandF()ra~in~ W<I luisE.AcostaandGlaucoMachado 9 NaturalEncmies 11f.J JamesC.CokendolpherandPlamenG.Mitov ]() Defense~lcchanisl11s 1/-1 PedroGnaspiniandMarcosR.Hara 11 SocialBehavior -Inn GlaucoMachadoandRogelioMacias-Ordonez 12 Reproduction n-l GlaucoMachadoandRogelioMacias-Ordonez 13 Development -I'is pedroGnaspini 14 Ecophysiology -171 FlavioH.Santos IS MethodsandTechniquesofStudy 4XY References 525 TaxonomicIndex 575 SubjectIndex ;X7 Contributors LuisE.Acosta GonealoCtrtbct CONICI."T- FacultaddeCienclasBxactas. DepartmentofOrgantsmtc&EvolutionaryBiology FtsicasyNaturales andMuseumofComparuttvcZoolol;:Y UniversidadNaclcnaldeCordoba HarvardUniversity Av.veleaSarsfleld299 16DtvtnuyAWllUC 5000Cordoba.Argentina Cambridge..\IA02118,USA SarahL.Boyer PedroGnaspiut DepartmentofOrganismic&EvolutionaryBiology IA..purtamcntodci',()olog,ia andMuseumofComparativeZoology lusututodcBtodcndas HarvardUniversity ttnlvcrsidndcdeSill)l'illllll,l']'11294 1hDivinityAvenue 05411-970SaoPaulo.SP.Brazil Cambridge.MA02138.lIS,\ liir~l'nGruber ThomasS.Briggs Nuturhfstortschcsxtuseumin\\'iL"n DepartmentofEntomology Hurg,ring.; CaliforniaAcademyofSciences ,\·1(1!4wlcn.Austrfu GoldenGatePark SailFrundsco.C,\9411S.USA 1\1.1I·('osH.llar;1 Dcpartumcutodei'J.KIIII~i;1 lamesC.Cokendolphcr tnstuutolIl'Billl:,iL;ndas NaturalScienceResearchLaboratory ltuiwrsidadc-deS,iol'uulo.CP1l2.lJ4 xtuscumofTCXilSTechUntvcrslty tl',411-970S{lol',mltl.Sl~Brazil Lubbock.TX7lJ-!(IY.(ISA ,\l!ri'lI1CJB,h:ur~ DavidJ.Curtis Ikp,lrtilllll'nl(}dl'lnwrtcbrudos DepartmentofBiologit"alSdt'lHX'S .\hISl'll;\adoll,d UntvcrsnyofParsley llnin..'rsidilliedoBrasil Scotland.UnitedKingdom tjumtudOlBoa\'isla 111\j4(J-0411Hiulit'[anciro.HI.Brazil Jason,\.Dunlop InstitutefilrSystcmuttschci'.llt)[tlgil' C;1"Ul'Ui\la('llal!ll MuseumfurNarurkundcdcrHumboldt j\lllSl'UdeIlbl(')ri,1.'alllnli lilliwrsiliilzuBerlln lnsuuuodcBillhl~i<l lnvalldcnstrussc-l1 llum-rsidack-E"'ladu<l1del·ampilla....~'I'hJill) D-I0II:;Berlin.Germany 1WSPJ,OCall1pina....Sl~Brazil I{II~diu\I,u'ia..-C)nli,ill'l l'cu-rSdl\\'l'ndingl'r 11q1;lr!;IIlIl'llIpdl'1:iplul-tl<ll:nlluli\';1 :\lllSl'llllldhlsrotn-nuruu-llc Iil..l;illlll1dl'I:I·ulul-tI;1 I,routetil'I\lala~llou,('f1-12I1 \.l',..\[1.1'1I"1;11h: 1:I'lIl'\"l'h,Swilzl'r1i1lul \.11.11"1.\IT<lITlll,"1111111,\kxko IdTn'~'\\',Shultz \1lIi1l1dil(',,\h'lIlll':O; lh'purfnu-ntofElll(lll1ol()~y III'parl,lllll'1l1l1dl'111\'I't"ll'hnldllS lIni\'l'rsilyof:\Iaryland \IU"l'U\ill'illll'.! ('lllk~l"Park.1\11)1(1,-1-2,lISA lum-r..i1Iol1kdlll:n.si! I.JuillWdilnOil\"i..ta ('hdsillphc'.rK,Ti.I}'lnr ..!119-l-II·tl-l-lll{illdl'[nnviro.R[,llrilzil DcpuruucruofTcrrcstrtallnwrtcbrurcs \\bll'fllAuxtrullunMuseum I'lillI1('11(;,:\lilm J.Od'I'l!Bag.49,\\'l'IShp(lll]hl)Sh lk'parlllll'lIlofi',lIolu:..:rand'\lIlhropulo~y \\'l'slt'rn,\uslralia,Australju Fcll"llhyorBioi,,:...):" UnlvcrsltvofSlllhl An..l.uci..Iourtuho .'il)ril~cIIlZ;ll1kl1\'ttlvd. INII,\- lnslilult)NactonuldcPcsqutsasdaAmazoutu tIh-l-Solia.Blll~ilria t'oordcnacaodel'csqutsusfA'ol6giras/CPEC·Sctor\'s AwmdaAndreAruu]o.19J6-Aldxo,CPol7S h9(1!1-9,11\Ianaus,A~l,Brazil ,\hl,ll'i'n'I.(;IIl\Ii'.lll'l I)l'parlclllll'lliodl'tnvcrtcbrudos NobuoTsurus..kl t.u\jluullttSvulct'UursNdlduuucdBlcoonadno\l'hBlraa~i1 FL'aIlb'uolrtaytoofryRocrgBtoiouialllgSYciences 211IJ-II-I-II-l-III{iodeluuctro.RJ.Brazil TTooluloorni.UhmMOw-Mrs5tt,y)I,Japan HkurdoPintu·da-Hudli1 DurrcllUblck lk'parlilllll'nlodei'.(I(I](~ia DepartmentofEntomology tustltutodeBi(ll:k'ndi.s CaliforniaAcademyofSciences llnircrsldudcdeSiloPaulo,0'I1-1-6I ColdenGatePark 0;;-1-22.·9,0SilOPaulo,Sl~Brazil SanFrancisco.CA9411X,USA I-'Ioh-inII,Silnlos RodrigoII,\\'iIIemart DepartamentodeZ()()It~ia Departamentode7.oologia InsnuuodeBiodl'ndas InstltutodeBiocienclas UmvcrsldadcdeSilOPaulo,CPI12lJ-I UnlvcrsidadcdeSuePaulo,CP11294 OS-l-22-9,OSilOPaulo,SP,Brazil 05422-970SilOPaulo,SI~Brazil Preface TheideaofwritingabookonOpilionesstartedmorethan[ouryearsagoasare suitofthe.needforacomprehensive\'OIUlllCthatencompassedallaspectsofhar vcstmanbiology.ashadbeendonepreviouslyformites.spiders.scorpions.pscu doscorplons.camelspiders.and whip spiders.Although harvestmen arc the third-largestgroupofarachnidsandoneofthemosrconunonfindsofamateurnat urallsts.Htttesyntheticworkouthclrbiologyandsystematicshadbeenputtogether. Thisbookserves thispurpose.andforthisreasonithasbeencompiledbyalargl' bodyofexpertsworkingin11countriesintheAmericas.Asia.Europe.andAus tralia.ThisisthefirsttimethatalargefractionofIhL'uvallublclltcrururctillhar vestmanbiologyandsysrcmattcs-c-Includlngworkswritteninmuuydtffcrcnt1(1)1. guages-chasbeenpUItogl.-thcrinacomprehensivevolume.Indoingso.theauthors hawreliedonstudiesoftheuniqueanrfbutcsofOptlfoncshi()lo~..:y,aswellasOilgcn era!behavioralandecologicallnvcsnguuonsconductedusinghurwsummSIXr'il'S Thishookshouldproviderbchucrnuuonaluudtcnccwithabroadiuxononucand ecologicalframeworkforunderstandingOpiliolll'Samicompunugthem withspi· dcrs.scorpions.andotherwell-studiedarachnids. BroadinSCOPi',thisvolumeisutmcdatansweringrelevantqUl'stiollSIromudi· vcrsltyoffields.whileillthesamcnmcimlkalingareasinwhichuddtnonulrcscurch isneeded.Itisfurthermoreintendedtuprovideunin-depthsununarvofourcurrent undcrstundtngofharvcsrmanbiologyandsystcmam-sbothforre-cun-hcrsaln-udy ucquumrcdwithharvestmen andfortlu-untnulutcdr0I11Il11lllily. 'l'hus thebook shouldbeofgreattutcrcsrIIIaruchuoloalsts.aswellas10:wologisIS.l'rol,,:...:isIS.ami amateurmuuralfstsingencml.TIll'rhupu-rswerewrittenwlthaninlntlbdplinar) audfcnccinmind.andtheygi\'l'broadsynthctk-vtcwswithintheirdiffcrcnrspcciul lit's:theresult.wehope.isrlnuthetextispalillabk'l'lIoughIIIuuracrrhcinh-n-st01 expertsand11111lLX'lx'rtsalike.includinggruduuu-students. ThehooklscomposcdofI::;chnprcrswrtucuhy2::;muhors.,\IlhllUglll'~ll'lISi\'l' editorialworkwasneeded tom'C0l11l110d,lteIlu-11l'lerogel1l'ityofsIrlesnnd10muid thetradinonalproblemsofahookwnhmanyouunluuors.ururcfu1n-mh-rmaywr taiulyencounterSOIlIl'iIlCOllsistl'nl'il'S.SOIlIl'lopkshawIxvnlouclwduponiumon thanonechapter.althoughwchuwattempted10elilllilwil'n-dundum-icxilS1I1tlCha... possible.Otherlopkshawnetbeendiscussedilldl'lililillIhis\ulullll'lx-cuusc01'IIll' lackofInformationuvatlnbk-illtlu-hurvcstnumlncnnurcorbo-auscrhcn-arc-cur rcntly110activeexperts011rhcsubkxt.Howewr;\H'arl.'l'lUllidl'tlllh;IIIIll'<!ualily.nul depthorIIll'mfomuunmprovidedinIIll'hookwillxuppluntnnv(ll'f;ISiollill.uu!..s.iull x I'll'I.l('\' l"'lllpilill;:'Ud1anllllprdwll,in'volumewasnOl'asytask.andthepron-sstook alm,"'l,',Iur.n"lr,.l)lIrill~Ihistinn-In'n-ccivrdtill'ln-lpofmunype-opletowhomwe arc~rl·;III.I-inth-htcd. Fir..tIll'ull.Ill'111<lIlkrhcouurlbutorstothisbook.whopu liclIlhde-altIIhhourmanyrcquv..rsandhelpedduringlhel'ditingprocess.Allchap rcr-,Ill'rl'lll·l'f-n'\'il'\\l'd.amithusweextenduuracknowledgmentsandgruutudcto allLhl'pn'fl...v.ionul-,\\htll'ardlillyrl'\·iSt'.dtill'd1apll'rsofIIll'nook.\\'l'fullyn-ulizc thut fl'\ie\\ill!!-muuu-crtpt-,boru'oftill'lIloslaltruistictilsksundtukcsenormous iIIlhlllllt""flillll'alldcfhu-t..ha<mulltokenofnurapprl'dalion.llll'numesofallrc Iklln..urcli..u-dlwrctill,lilutil1l1Sl1rcontrlbutorsurcgi\'l'llinthescrtlon"Coutrih 1I111!,'''):l.ui-,I:..\co..ru.DultoulIl'Sollza.uuortmttutvcrsldadcdcSilOl'nulo.Brazill. \Iurra~S.lthnurtmvcr..ilyofl;l'orgiilru.uhcns.I"S..\I.SarahBoyer.AfunB.('<ldy l\li,lluilniwr,ity.IS.\I.lonurhuu.\.CIlll.dillgtnnt\"Cltiollal.\hISl'UlllofXuturul lIi'hlr~-and Srnilh,olliilnlnsnuulon.I'S.\I. lamesC.Cokcndolphcr.Richard\1. 1>uflidd1I111\\imllnin'r,ity.I'S.\1.\\"illiillllt;.Hx-rhardtttnlvcrstdadde('oslaRicu andSIlIilhsnni,II1lnxlitution.Co,1ilHiruI..\\;Ik'ollllEdmundstllntvcrsityofCcntrul lnucashin-.tnucdKmgdonu..\1<111'.:.\.ElgarII'l1in-rsilyof.\ldbollfl1l',r\ustralial, \larn'I'1(l.t;IIIlZilgall·lliwrsidalkE...t.adualdeCamptnas.Brazill.PhilltpeGrund l'llla,t.\lU-.iU'lll:\iltinnald'llishlirl'Xururcllc.l'raucct.[urgeuGruber.RobertHolm tll'r~L\lh<lhi!-.l·ilIniwr,ity.Calladal.Taplx'y11.Jonl'Sl\'irginia.\Iiliturylnsunuc. IS.\I,.\drianllB.Kur~:[ohnILB.LightonIt'niwrsilyofXcvada.lISAI,Rogcho \laci,I'-()nkll1l'z,.\lich<ll'l.\k,\1OI11l111IliwrsilyofCOl1l1l'clil'ul.IISAl,FernandoL.I~ \larqlll'stI'nin-rsidtldl'deSilOl'aulo.Brazill.[oclu-u.\!tlrll'nsrlnsttnnfurZootOg.il'. \Iaim.l~l'nnallYI..AdrtunoS..\IdortntvcrsldadcFederaldoRioGrandedoSuI. Bl"iJzill. Dtlllgiass II. \lorSt'.tBrown 11niwrsity. t"S..\I. ..\rturo .\Iuiloz-('ue\·as t.\IU"':'Ulll'aliollalll"lli,loirl'Xatun-llc.Fruncct.ToneXovakIFacohadiPedagogta. SloH'niill..\Ild I\;rl'zt;ollzi.k,z.tcorgc:\.PotnurtOn'gullStateUnivcrslty,lISAI. lcronwC.Ikgil'rII'ni\'l'rsityorxtnrvfund.I'S:\I.Adalbertolosl'dosSantosttnstltuto BlI{<Jnl,1J1.Brazill.DavidS.SaunderslIinin-rsityofEdinburgh.Scottandr.Peter Sdl\\'l'lldingl'r.PaulSeldenll'niwrsilyofxlunchcstcr.UnttcdKingdomI,\Villi<lmA. ShearLllallllx!l'n-SydncyColh-gc.I"S.-\I.Jell"rl'y\\:Shultz,DouglasW.Tallamy(llm wrsityofDelaware.I"SAI.XobuoTsurusakl.DarrellIlbirk.Peter\\'ergoldtIAlhert LudwigsI-niwrsiliil.t~l'rnlilllyi.and\'itoZingl'r1l'l:"allifillHistoryfl.tusl'umSoulh TyroJ.llalYI.Finally,\\'l'wouldlikl'tothankRonClouseandlimilesson.whokindly l"l'\'isl,dIIll'nmll'ntsandEnglishgrammarofSl'WfilldUlpll'rs,Humbl'r\oYamaguli furIwlpingwilhthl'edilionoftill'plilles.andAnnUowncr-HazcllfromHarvardUni \"l'rsityI'rt'Ssforslmringhereditingexpertiseilndpatientlydealingwithourmany n't:Jlll'SlsandSdll'dulcchangl'S.Thisbookisdl.>dicall'd10ourfamiliesandsludcnts fortlll'irunderstandingandsupport.11isalsodedicatedtothreeofthegreatest modl'rnopilionolog.ists.ChristianJuoerlhic:,Jochl.'nMartens.andf\l.ariaRambin, \\"ll()S'linddatigable\\'orksl'n'cdasaninspirationtous.
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