Sociobiology PDF
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Sociobiology THE ABRIDGEDE DITION EdwardO .W ilson Drawbyi Snagrsa h Landry Sociobiology THEA BRIDGEEDD ITION Byt heS ameA uthor OnH umanN ature Castaen dE cologiynt heS ociaIln sects witGhe orFg.eO ster SociobiolTohgeyN :e w Synthesis Lifoen E arth withT homaEsi sneWri,n sloRw. BriRgigcsh,a Er.d DickerRsoobne,Lr .Mt e tzenbReircgh,aD r.Od ' BriMeinl, larSdu smaWni,l liam E. Boggs TheI nseScotc ieties A Primeorf P opulatiBoino logy witWhi lliHa.Bm o ssert TheT heoryo fI slanBdi ogeography witRho beHr.t MacArthur Sociobiology THEA BRIDGEEDD ITION EdwardO . Wilson THE BELKNAPP RESS OF HARVARD UNIVERSITPYR ESS CambridgeM,a ssachusetts andL ondonE,n gland Copyrig©h t1 97159,80b y t heP resideantn dF ellows ofH arvarCdo llege Allr ighretsse rved Printientd h eUn iteSdt atoefsA merica Sevenptrhi nt1i9n9g8, LibroafCry o ngCraetsasl ionPg uibnlgi Dcaattai on Wilson,E dwarOds borne1, 929- Sociobiology. Bibliograpp.h y: Includiensd ex. 1.S ociable haviionra nimalsI.. T itle. QL775.W553991 .57 9-17387 ISBN0 -674-81623-4 ISBN0 -674-p8b1k.6 24-2 v Prefatcote h eA bridgEeddi tion Modems ociobiiosbl eoignycg r eabtyeg di ftiend� uatset udeinsta st ,e stamteotn htce o mmunaolfi ty vestigwahtoow rosr pkr imariinpl oyp ulabtiiooln science. ogyi,n vertebrate zooleongtyo,m oilnocglyu Tdhienssgec ientailssctors i tirceaalmdlo ys ot ft he especiaanldvl eyr,t ebzroaotleoB geyc.a umsyet rainchaptienre sa rdlrya fIta .m especigarlaltyet fou l inagn dr eseaerxcphe riweenrcfeeo rtuitiontu hsel y RobeLr.tT rivfeorrrse adimnogso tf t hbeo oakn d firtswtos ubjeacntdts h ewraess ommeo mentluemf t discusiswtii ntmghe f rotmh tei moefi tcso nception. frowmr itiTnhgeI nseScotc ieItd ieecsi,dt eold e arn Otherwsh or eviewpeodr tioofnt sh em anuscript, enougahb ouvte rtebtroaa ttetse mapg te nesruaml witthh ec haptneurm belriss taefdtt ehre niarm es, maryT.h er esuwlatsS ocioby:iT ohlNeoe gwS ynthesisa,r Iev aCnh as(e1 3I)rv,e Dne Vor(e2 6J)o,hF n.E isen publisihn1e 9d7 T5h.e b oomke tw itshu bstantibaelr (g2 223,2, 4 2,5 R)i,c haDr.Ed s te(s2 3R)o,b ert critsiuccacleI sths a.sb eewni deulsye tdh roughouFta ge(n1 -57,) M,a dhaGva dg(i1l- 5R)o,b eAr.t thweo rladsa s ourbcoeo ikn r eseaarnca hd,v ancedH ind(e7 )B,e rHto lldo(b8l-e1rF3 .)C ,l arHko well textbaonodakg ,e nerreafle rweonrckHe.o weveirt,s (26S)a,r aBhl afHfredry( 1-1135,- 1266,) A,l ison larsgiez( e6 9d7o uble-cpoalguemasnn) dn ecessariJloyl (l2y5 A.)R ,o sKsi es(t7e1r,1 -1B3r)u,c Re.L evin higcho sptr evenittfe rdo rme achmiuncgh o ft he (45,)P,e tRe.rM arl(e7rE) r,n Msaty (r1 1-1D3o)n,a ld laragued ieonflc aery e adaenrdss tudewnhtosh ave W.P fa(f1f1 K)a,t herRianl(el1 s5 J)o,Sn e ge(1r- 86-, becomien tereisnt esdo cioabnidio nlt ohmgeay n ner 132,6)W,.J ohSnm it(h8 -1R0o)b,e Mr.tW oollacott inw hicthh ipsa rticwuolrfakor r muliattG.ee dn eral( 18J)a,m eWse inr(i1c-h85 -,1 3a)n,dA motZza havi readewresrf eu rtdhiesrc ourbaygt ehdte h ickoeft s (5). tecr.ih.cadli scussainoddn ast sau mmariIents h.e Illustrations, unpubalnidst heecdh manuscripts, presevnetr siSoonc,i obyi:To hleAo bgridEgdeidt ionn,i caadlv iwceer seu pplbiyeR d.D .A lexanSd.Ae .r , Ih avter immtehdte e xdto wnt oi tess senitnitarlo BoormaJna,c Bkr adbuFr.yH ,. B ronsoWn., L . ductoprayr tasn dm ositn terecsatsiheni gs torieBsr,o wnN,o amC homskPy.,A . Comingl,a in whilree taitnhifenu gbl als sitcr ucotftu hroeer iginaDlo uglas-HaMmairlJyta onnWe,e sEtb erhaJrodh,n bookT.h issh ortevneerds iiosin n tentdoes de rve F.E isenbRe.rD g.,E steCsh,a rlGeasl tV,a lerius botahs a textbaonodka semi-popguelnaerra acl GeisPte,t eHra asW,. J.H amiltIoInIB ,e rHto ll counotf s ociobioBleocgayu.os fet heu nusual doblSearr,aH hrd yA,l isJoonl Jl.Hy .,K aufmaMn.n , amounotfi ntearnedsc to mmentiahtra ygs e neratedH,.A .K eenleyAs.iR d.Ke i,e stHearn,Ks u mmeJr,. Ih avlee tfhtfe i ncahla potneh ru masno cibaelh aviorA .K urlaMn.dR ,.L eiBn.,R .L eviPn.R, . L eviPt.t , (Chap2t6ei rnt haeb ridvgeerds ivoinr)t uianltlayc t.R .M arlEernrs,Mt a yrG,.M .M cKayD,.B .M eans, Inw ritionngg e nersaocli obiIoh laovgheya ,dt o A.J .M eyerriMeacrktsMi,on y nihRa.nA ,.P aynter, ventuirnetm oa nyd iscipilnwi hniecsIhh avneo d i Jr,K. atherRianlelL sy,n Rni ddifPo.Sr .dR ,o dman, recetx periaenndct eh,uh sa dt or eloynt heex pertisLe. RLo.g eTrhse,l Em.aR oweWl.lE ,. S chevNi.Gl .l , anda dviocfce o lleaagtHu aersv aUrndi verasnidt y SmitJhu,d Ay. S tampRs.L, .T riveJr.Ws .T, r uman, elsewhTehreeg .e nerotshietdyyi splagyueidd,i ngJ aWni ndE,.N .W ilmseEn.E, .W illiaamnsdD, .S . mep atietnhtrloyuf gihla mnsdp ublicactoirornesc,t Wilson. inmgy e rroarnsdo, f fertihnkegi nodf e nthusiastiKca thlMe.e nH ortoans sisctleods ienlb yi blio encouraguesmueanrltel sye rvfeodpr r omisgirnagd - graphriecs earcchhe,c kmeadn yt echndiectaali ls, vi Prefatcoet heA bridgEeddi tion andt ypetdh em anuscrtihprto utgwho i ntricatoefm ya rtiicnBl uelsl oeftt ihEnen tomolSoogciicoeaftl y drafNtasn.c Cyl emenetdei ttehdme a nuscrpirpot, Ameri(cvao1 l9.p, p .2 0-2129,7 3S)c;i e(nvcoe1l 7.9 , vidimnagn yh elpsfuuglg estcioonncse ronrignagn ip.4 661,9 7c3o;p yri©g 1h9t6 199,7 b3y,t hAem eri zatiaonnde xposition. canA ssociaftoitroh neA dvancemoefSn cti ence); SaraLha nderxye cutthededr awionfga sn imsaol ScienAtmiefirci (cvaon2l 2.7p ,p 5.3 -5149,7 2C)h;e mi cietpireess enitnCe hda pt1e9r-s2 I6n.t hcea soef caElc olyo (gES.o ndheiamnedJr .B .S imeoendes,,. thvee rtebsrpaetceih eesrc, o mposiatiraoemn osn g AcademPirce s1s9,7 0M)a;na ndB easCto:m parative thfier sttor epreseennttis roec ietiinte hsec,o rrectS ociBaelh av0i.oF r.E isenbaenrdg S.D illon, W. demograpprhoipco rtwiiotnahssm , a nys ociianlt ereds,S. mithsoInnisatni tPurteis1os9n,7 0T)h.eq uo actidoinssp laaysce adpn l ausbiebi lnyc luidnoe nde tatifornostm h eB hagavada-rGteia tkaef nr otmh e scenIeno. r detorm aket hder awiansga sc curaast e PetPearu pPerre tsrsa nslTahteie odni.ta onrdps u b possiwbels eo,u gahntdw ergee neroguisvletynh e lishaerrtesh ankfeotdrh epierr mistsoir oenp roduce helopft hfeo llowbiinogl ogwihsoht asdc onductedt heesxec erpts. reseaornct hh seo ciobiooflt ohgey indisvpiedualI w isfhu rtthoet rh antkh feo llowaignegnc aineds cieRso:b eTr.Bt a kk(erre constorfut chatepi poena rindivifdouprae lrsm istsoir oenp rodmuactee rfioarl s ancaen dp ossisbolcei baelh avioofdr i nosaursw)h,i cthh ehyo ltdh ec opyriAgmhetr:i cAasns ocia BriaBne rtr(almi olnasi)Dn,o uglas-Ha(mAiflrtiont iofno trh eA dvancemoefSn cti enrceep,r esenting cane lephaRnitcsh)a,Dr .Ed s t(ewsi dlodg sw,i lde ScienAcmeer;i can ZAononluoRageliv site;Iw ns;c,E . . beesFt.C) l,a Hrokw e(lrle constroufpc rtiimointsi vJe.B riClol;. C ambriUdngiev erPsrietsyAs m;e rican mana ndt hPel eistmoacmemnaefl a unAal)i,s Joonl ly SocioeftI yc hthyolaongdHi esrtpse tolroegpirsets, (ring-tlaeimluerJdsa )m,e Msa lco(lwmi lddo gs), sentCionpge Dioaw;d eHnu,t chinasnodRn o sIsn,;c . JohnH .K aufma(nwnh ip-twaaillleadb Hiaenss), DukeU niverPsrietasyns d t hEec ologSioccailoe ft y Kumme(rh amadrbyaabso onGse)o,r gBe.S challerA merirceap,r eseEnctoilyno;Dgg r.M arJya nWee st (gorilalnadGs l)e,En . W oolfen(dFelno rsicdrau b EberhaWr.d H;. F reemaannd C ompanrye,p re jaysE)l.sS o.B arghoLoerns,l Ai.Ge a raayn,dR olla sentSicnige nAtmiefirci cDarCn.h; a rlSe.Hs e nrDyr;. Tryoand deadd viocnet hdee pictoifto hnse u r J.A .R .A .M .v anH oofHfo;u ghtMoinf flCionm M. roundivnegg etaOttihoendr.r awinignts h ibso ok panyM;a cmilPluabnl ishing CIonm;cPp .raonfye,s wereex ecubtyeJ do shBu.aC larakn,dm osotf t he sorP eteMra rleMrc;G raw-HBiololkC ompany; grapahnsdd iagrbaymW si lliGa.Mm i nty. Methuaennd C o,.L td;P. rofesDsaonriO etlt eD;r . Certapians sahgaevsbe e etna kewni tlhi totrnl oe KatherRianleRl asn;d oHmo usIen,;c P .rofeCsasrolr change TfhreIo nms eScotc iebtyiE e.Os ., Wilson W.R ettenmtehyReeo ry;aS lo cieLtoyn,d oUnn;i ver (BelkPnraepso sfH arvarUdn iverPsrietsy1s 9,7 1); sitoyfC alifoPrnrieasT sh;eU niverosfiC thyi cago theisnec lsuhdoepr otr tioofCn hsa pt1e,3r s,6 ,8 ,9 , Presisn,c ludrienpgr esenotfTa htAeim oenr iNcaatn 14a,n d1 6i nt hper esebnotoa ks w elalsa s ub uralainsdWt o;r tPhu blisIhnecr.s , 13, stanptoiratloi foC nh apt1e9wr,h icphr eseanb trsi ef Finalmluyc,ho fm yp ersorneasle arrecpho ritne d revioeftw h seo ciinasle Octthse.er x cerhpatvbsee en theb oohka sb eesnu pporctoendt'i nuboyut shley takefnr oAm P rimoefPr o pulaBtiioolnob gyEy .,O . NatioSncaile nFcoeu ndatIitio sfn a.i tros ayt haIt \Vilsaonnd H.B oss(eSritn auer As1s9o7c1i)a,t weosu,ln do hta vree achtehpdeo ifnrto wmh icahs yn W. LifoenE artbhyE, O. . W ilseotan l (.S inaAuser thesciosu lbdea ttempitfie thd a dn obte efno trh is and socia1t9e7sO3,t) h.e pro rticoonmsfe r ommy a rticleg enerpouubsl siucp port. '/GroSuepl ecatnidoI ntS si gnificfaonErcc eo logy" (BioScvioeln2.c3p e,p, .6 31-613987,3c )o,p yri©g ht E.O.W . byt hPer esidaenndFt e lloowfHs a rvarCdo l CambridMgaes,s achusetts 1973 Othepra ssahgaevsbe e eand aptferdov ma riousS eptem1b9e7r9 lege. vii Contents PartI SociEavlo luti1o n PartI I SociMaelc hanism6s5 1 TheM oraliotfyt heG ene 3 6 GroupS izaen dR eproducti6o7n TheD eterminoafGn rtosu Spi z6e8 2 ElementaCroyn ceptosfS ociobiology 7 AdjsutaGbrloeu Spi z7e1 BasDiecif niti7o ns TheM ultiplaincdaR teicoonn stiotfu tion TheM ultiEpflfiee9cr t Socie7t2i es TheE volutiPoancaermyaa knedSr o ciDrailf 1t0 TheC onceopfAt d aptDievmeo grap1h1 y 7 TheD evelopmeanntd M odificatoifo n QualiotfSi oecsi a1l2i ty Social Beha7v4i or TheC onceopfBt e havioral1 4S caling TheD ualiotfEi veosl utiBoinooaglry1y 5 TracktihnEegn vironwmietEnhvt o lutionary Chang7e5 TheH ierayor fcO hrgnaismic Res7p6o nses 3 TheP rimeM overosf S ociaElvo lutio2n0 TracktihnEegn vironwmietMnhot r phogenetic PhylogeInneetrit2c0i a Chang7e7 EcoloPgriecsasl2u 3r e HormonaensdB ehavi7o7 r TheR eversiobfSi olciiEtavylo lut3i1o n Learnin7g9 Sociali8z1a tion Play 4 TheR elevaPnrti ncipolfeP so pulation 84 TradointC,iu ltuarnedI, n vent8i6o n Biolog3y2 Microevol3u2 tion PhenodevainadGn entest Aiscs ilmaiti3o4n 8 CommunicatiBoans:i Pcr incipl9e0s InbreeadnidKn ign sh3i5p Humavne rsAunsi mCaolm munica9t0i on AsosrtaatnidDv ie satsastoMiravtei n3g9 DiscrveetresG ursa dSeidg na9l2s PopulaGtrioownt3 h9 SignSapclei fic9i3t y DensDietpye nde4n1c e SginaElc onom9y4 Intercompe4n3s ation TheI ncreoafIs nef orma9t4i on PopulaCtyicolonefM s a mmal44s LifTea bl44e s TheS tabAlgeeD istrib4u6t ion 9 CommunicatiFounn:c tioannsd C omplex ReproduEcftfiov4re6t Systems1 00 ranKd Select4i7o n TheF unctoifoC nosm munica1t0i0o n ComplSeyxs te1m0s7 5 GroupS electainodnA ltruis5m0 GrouSpe lect5i0 on 10 CommunicatiOorni:g ins and Interd(eImnitce rpopSUellaetcit5oi1no )n KinS elect5i5o n Evolutio1n1 0 Reciprocal 5A8l truism TheS esnorCyh anne1l1s3 AltruBieshtaivci5 o9 r EvolutiCoonmapiretyti aomno nSge nsyo r TheF ieolfRd i ghotuesne6s2s Channe1l1s6 viii Contents 11 Aggressio1n1 8 18 TheC oloniIanlv ertebra1t8e3s AggresasnidCo onm piettio1n1 9 TheA daptBiavseoi fCs o lonia1l84i ty TheM echaniosfCm osm peti1t2i0o n GeneErvaoll utiTorneanirdnyCs o lonia1l8i5 ty TheL imiotfAs g gsrseoin 121 TheC oelente1r8a6t es TheP roxiCmaatueso efAs g gsriseon1 22 TheE ctopr1o8c7t s HumaAng gress1i2o6n 19 TheS ociIanls ect1s8 9 12 SociSapla cinIgn,c ludiTnegr rito1r2y8 WhaItsa S ociIanls ec1t89? A "TypiTcearlr"i tSopreicai1le2 s9 TheO rganizoafIt nisoteSn co cie1t9i1e s TheM ultiFpolremo sfT errit1o2r9y TheP rimMeo veorfsH ighSeorc iEavlo lution TheT heoorfTy e rritEovroilault1 i3o2n inI nsec1t93s SpecPiraolp erotfTi eersr it1o3r4y TheS ociWaals p1s9 6 TerritaonrdPi oepsu laRteigounl a1t3i6o n TheA nts1 99 TheS ocBiaele 2s0 1 TheT ermit2e0s3 13 DominancSey stems1 37 ExampolfDe osm inaOnrcdee r1s3 8 20 TheC old-BlooVdeerdt ebrates 207 SpecPiraolp erotfDi oemsi naOnrcdee r1s4 0 TheA dvantoafgB eesi Dnogm ina1n4t1 FisShc hoo2l0s7 TheC ompensaotfBi eoinSnsug b ordi1n4a2t e TheS ociBaelh avoifFo rro g2s0 9 TheD eterminoafDn otmsi nan1c4e3 TheS ociBaelh avoifRo erp ti2l1e2s IntergDroomuipn an1c4e4 ScaliinAn ggg resBseihvaev i1o44r 21 TheB irds2 14 TheE voluotfiC oono perBarteievdei 2n1g5 :,;, Roleasn dC astes 146 TheJ ay2s1 8 TheA daptSiivgein ciafnocfRe o le1s4 7 TheO ptimizoafCt aiesoS tny ste1m4s8 22 EvolutionTarreyn dwsi thitnh e RoliensV ertebSroiacettei 1e5s0 Mammals 220 RoliensH umaSno iceti1e5s4 GEmerPaalt te2rn2s1 TheW hiptWaailll a(bMya crpoapryruis)2 2 4 Dolphi2n2s5 15 Sexa ndS ociet1y5 5 TheM eanionfSg e x1 56 Evoluotfit ohSnee Rxa ti1o5 6 23 TheU ngulataensd E lephant2s3 0 SexuSaell ect1i5o8n Chevro(tTariangsu l2i3d1a e) TheT heoorfPy a renItnavle stm1e62n t TheV icu(nVai cvuigcnaua)g 2 n34 TheO rigoifnP so lygaMmoyn,o gaamnydP, a ir TheB luWei dlebe(eCsot nnoecst haianueur)ts 235 Bondi1n6g4 TheA friEclaenp h(aLnotx oadfioarcnntaa2) 4 0 Communal Di1s6p5l ays OtheUrl timCaatueso efSs e xuDailm orph1i6s6m 24 TheC arnivore2s4 2 TheB laBceka (rrU sausme arni)uc s243 16 ParentCaalr e 168 TheL io(nPn ahtearl e)o 243 TheE coloofgP ya renCtaarle1 68 WolvaensdA friWcialnDd o g(sC anid2a4e6) Parent-OfCfosnpflri1ic7nt0g ParenCtaaralen dS ocial Eivnot lhuest eiIcontns 25 TheN onhumanP rimate2s5 1 and Mamm1a7l3s TheD istinctiTvrea ioStfPos rc iimaalt2 e51s AlloparCeanrte1a 7l4 TheE coloofgS yo ciBaelh aviinPo rri mat25e4s Adopti1o7n5 TheL essMeoru sLee mu(rM ioccrembuuisrun s)25 6 TheO rang-(uPtoannpg yog ma)e 2u5s7 TheW hite-HaGnibdbeod(n Hy lotbelasa r2)5 8 TheR ing-TLaeimlue(rdL emcuart) t 2a58 177 PartI IIT heS ociSaple cies TheH amadryBaasb o(oPna hpaiomr ayd)a s259 TheE aesrntM ountGaoirni (lGloar giolrlial la 17 TheF ourP innacloefSs o cial bereiin2)g6 4 Evolutio1n7 9 TheC himpan(zPeatenr ogtle)os 2d6y5 Contents ix 26 Man:F romS ociobioltooSg oyc iolog2y7 1 Territoanrdi Tarliibtayl2 i8s9m PlastoifSc oictiOyar lg aniza2t7i2o n EarSloyc ial Ev2o9l0u tion BartaenrdR eciprAolctaruli s2m7 5 Later Social2 96E volution BondiSnegxa,,n dD ivisoifLo anb o2r7 7 TheF utur299e RolPel ayainndPg o lyeth2i7s8m Communica2t8i0o n CultuRriet,ua anldR, e lig2i8o4n Glossar3y0 5 Ethi2c8s7 Bibliograp3h2y7 Estheti28c8s Index3 49
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