Harvestmen: The Biology of Opiliones

607 Pages·2007·89.581 MB·English

by  Ricardo Pinto-da-Rocha, Glauco Machado, Gonzalo Giribet

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.