SEVENTH EDITION AN EVOLUTIONARY APPROACH A ARIZONA STATE UNIVERSITY SINAUER ASSOCIATES, INC. SUNDERLAND, MASSACHUSETTS TM C_ A ~ itIk ('OIlwy1' ~ informatioo. about .... fiP1~uponry 10 ri,'~ INln. ~aphby Ilruc'l' lyan. 1M FrontiJPi«~ 1lw lhrNt d;,.pa.y of tIw l-WnadJyas Nboon noion many qun"'- for n 'olutionary ~ ~aph by C K. 1.oftnz- Animal ~hovior.An Evolutionary Approach,~nfh Edifion C Copyright 2001 bySiN..... AS6Oriare.. lee. All rillh tll ........".<"d. This book may"''' be ...prod lK't'd in who'" <>f ill pan I.... any purp..... whal"'·.... without wriUffi f"-'rmission from the publisher. For inKmnaotion or 10 ord..r. odd""",: S inou~r A."..";,,t,..._Inc., P.O. 8.." 407, Sund..rlond , M.s..,,,hu..,Us. 01 375-0407 U.S.A. Fa. , 4 1 ].. .~ !l- 1lI 11 . IIll~mct ; publi. h...iN.....r.com; http://wv.w.•iN u~r.('<>m Akod, John, IQ.l2- Anim.a\ bfoty"ior / John Akork.-7lh <"d. r·mI. Indudor.. bobl"w'aphiuoI ...-ft,,,,, la'S. 1S8~ 0-8",,3-011-6 I. An,m.a\ behoo\'ior-E..olutioo.. I. Titl<- QL751 .A5Il 2001 591.5-.k21 lO'l H7 65 43 21 To Tht Naturt COl1st rvancy Contents in Brief 1 An Evolutionary Approach To Anima/Behavior J 2 Proximate and Ultimate Causes ofBehavior:How and Why BirdsSing 22 3 The DevelopmentofBehavior: A Focus on Heredity 50 4 The DevelopmentofBehavior:AFocus on the Environment 76 5 The Control ofBehavior; Neural Mechanisms 1/4 6 The Organization ofBehavior: Neurons and Hormones 154 7 Adaptationand Antipredator Behavior /82 8 The EvolutionofFeeding Behavior 214 9 Choosing Where to Live 248 10 The Evolution ofCommunication 280 11 The Evolution of Reproductive Behavior J 1f> 12 The Evolution ofMating Systems 300 13 The Evolution of Parental Care 394 14 The EvolutionofSocialBehavior 420 15 The Evolution ofHuman Behavior 456 Contents 1 I An Evolutionary Approach To Animal Behavior 1 Qul'Stions about Behavinr 2 II.,U' Qurs/ions nh",t Proximatf QWSfS 3 Wh!f Qut'ShOlls nwut U/h"",tr (nuSt'S 3 Ans",·..ring Pru~ima t<' and Ulhm.ote Qut.."tiOll5 abo", Bt'h.wior :; 8tn..:Jll'rS n..J H""""1l Btftn.ivr 5 Gulls and Egphrl/ R,."mt",1 9 Da rw iniolfl Theory and Ulti mate Hypoth......" 11 v..n.t,jnWtl Logic and 1MStudyof &Nwim 13 TM Probkm ""tlr Group sn~f;"" 14 T....ling AltemahH' Hypot~ 16 (Lm,i"'y,,,,1i $rim« 18 2 I Proximate and UltimateCauses of Behavior: How andWhy BirdsSing 22 D1ffnmt Songs: Proxim.oloPQ..-; 2.- s..o.d Uprnrna"'" Softg Dr,..Iop".", 26 nr AlIi.Pl Soflg C",,'rol S~..... 28 nr DtwIopmtntof''''' Sortg Conlrol 5~..... JI Sorr,~ Drffrrmcrs Bt1,.......,,,, So-uo Jl Awiding" MisunJm;tlllldill!i 34 Oifftonont Song$.: UltiINIe c.ll'O.'S J.I 1JIrAdh.villf 65 Difrermt SelKtion ",""UlYt'. Diffnnlt Centos. Diffnnll8eh.lvion 67 4 I The Development of Behavior:A Focuson the Environment 76 T1w InIWKti\'e' T1wofy of Dr\-eklpment: IiorrtIorll'!o .lind fltoh.o,ior 78 W/ItIl tol!«tIrIIt'.I FCItIl~ irt • HoNy 8ft' Colony so W/ItIl to~. Tm'itor¥/ ~ 82 E"~.nd Bt-hil"iof.1 Dr\'elnpmmt ll..f u rly Expt'Tietlc~ ,m4 ~Ilioll of RtlatilYS 85 I.tIIrm"8 as Bthal'wnll Ott",I"!'me"t 90 holution and fltoh.o,ior.ll Dr\-dopment 92 Thr ~ino V.t..,.o/ Dt(lflopll..~t"" f/nibility 94 Sn Dljfrre't1a5 ;rt S,..tWllNtrtillK AkJity 98 Thr EroIuhorr ofIwocioftit", u..mirtK 99 Thr Erolu/ioll of Ott",I",, """'t~t H~ts15i5 10j Thr Ada/'tit'" Value vf Dn>t1"P.......t,,1 Homro>lll$i$ 106 5 I The Control of Behavior: Neural Mechanisms "4 How Sn1n:Jm Cootrol BeN\v 116 StnMW)'~.,....rIJ 5..rrm./ J19 ~Ilg ..w ~illg '" Stn50ry /I'pIIl U6 UIIt" P.t"", GmmIIon US TIlt s..tg ofIW Mid.NI5drt>iwks 159 H_ DI10 Circ:adillll MKIloIllislllS WorA:? 161 w..g·TmnCydncfBml..... 165 T~ Ph!f.'iC/2/ EIII"rollmt'IJl llIfl"fflCf'i; LoIIg-TmII Cydl:S 167 Changing Prioritit-s in Changing Sucial Erwironl1>l"Tlb 170 TIlt Effrr:ts of u,n.l.litwI iii Iitt' C...... Anok 171 H~tklp~i:.tSocMlBoIMI'" 17J 71Adaptation andAntipredator Behavior 182 T1w MNlning of Adapl.llioo 184 T1w Comp.uati\·" Method for T..,;ting Adapl.lliooist Hypotlll"Sol."S 187 ~ Impvrlana of Dit\'rgffll ft\liw'iQII 189 TIlt Imp"',an.:t uf CoIl""":m ofOpti-t,ty 17wvry 136 TIttEroIllhoft '"Al/mW11A" OW/!; 136 Conwming "'hal You SeI«t lOEat 238 H_ to CJI;1tIt. Wlwlk 240 Why DI10 HII""'IU eoo..l1..... Akohol. 5Fn. Il1ld Dirt? 141 9 I Choosing Where to Live 248 H..bitat Selection 250 HiIblwt P'r,Ii"'iUS "'" Ttrntorilfl Sj;wlr5 ill Drf a Sign0' History of .. Sign.al-Rf'mving M..dwusm 2'l7 n", History of1n"d Wing. 289 s"n.1al Rt'Ui ,~n)( J07 Ricrivtrs Mily RI'1j~il'tH"IUS/ Sigrwls JOIj Why ~ lktJ'Iio>r OfyujSn Di/frrmtYo ill s,,'ual 5ell"Clioo and Competition for Malt'S 3rl SOCUZ/ Do...i""nct' and Millt FillUSs .HI Altl"l"lllllivr Milling TiKtics 332 A e-'i~ Siraltfl/ll'ifil AllnnaliR' Molti,,! TiICficJ J.U J1rI'tl' DrsIi>od Stnrtrgin, J1rI'tl'MoItlflg r.-t'" liS SavM smrtio>r I/ptJ 5"...... Coo'Ipt1" .... JJ7 Malt CWlrdin)( JJ8 s", u..l s"lt'ction ..nd Mat.. Choice 341 f~ .MIlt Chc>ict' ,,"11101<1 M11lmo1l &nqi15 J.I6 Ttstirrg IIwHnIlflry .MII~. Good Gnrn. ..>04 R~""""Y ~~J5J Sex......l Conflict l;oen.·een !\.tales ..nd Fem.l~ 353 12 j The Evolution of Mating Systems 360 Is Monogamy Ad..pti,·..? 362 M.mllllamy III .MI......../. J64 M"nllllamy i" Bird. J66 ulrl/-Pai, C"P"iIIfio>r.: n",.MIk Prrsp«fi.". 369 £xItI-Pllir CDJ'IUIIfiorr£ TJw FtrItIIh P....ptdit'" 370 Poly.andry without Pol)·gyny m Pol)·gyny J77 f"""/t [)o/nst Polygyny: Tht f"""lt Prnp«tiror 379 Rt50wIlY Dtfr- PolygyrIy J80 RrsouIlY [)%tst Polygyny, n", f(mak Prnp«riw JSl Saamlrlt Cc>mprlilion 1'oIYSY"JI JS2 Uk PoI!fSY'IY J85 Yo1ry Oct,\.Wts Aggrtpft;.. Ltb' J86 Why Oct MIIny F~ M111~ ll'illl flw Sa..... /o.Wks 1/1 Ub? JS9 I X T. l>I~ 01Ce>nlent. ~ ,13) The Evolution ofParental Care 394 , Why Is Parenl,ll Care More Often Maternal than Paternal? 396 EXCl7'tion~ to tlw Ruit 397 Wh .~ Do Mille Wattr Bugs Do All tlw Work? 400 Discriminating Parental Cart' 402 Offip"ng Rt>:ognition; C"",,,,,rati,,,, St udi<'s 403 Why Adopt Gmetic Strangtrs~ 404 TIlt History ofIntersptfi/ic Brood Parasitism 406 Why Accept a Parasite's Egg~ 408 Can Adoption &neftt fostl'l' Parmt.? 410 The Evolution of Pan'lltal Favoritism 411 • 14 The Evolurion ofSocial Behavior 420 The Bent'fits and Costs of Social Life 422 1be E\'olulion of Helpful Behavior 427 R«iprocal Altruism ar Pl'I'sonal GIlin? 429 Altrl/ism and Indim:t Sd«tim, 433 TIwAlarm Cail of Belding's Ground Squirrel 434 The Concept of Inclusive Filness 435 Inclu,iPf Fitn,,, and Helpers at tilt Nest 437 Insect Helpers al tilt Nest 442 1be Evolution of Eusocial Behavior 445 Testing tilt Haplodiploid Hypotht<;,s 447 Vrry Clll:'( Relattdnr5S I., Not E"-dition 01 Animal Beha,,;")]" dppe.>n'd. Purin!\ thi, time, much thai L, d"Preso;ing hils occurn.'<' polilk;, but i""tt'ad ronsider the pooitive side of the It'dl;t"f,which includes the many exce]- \tonI P"J""T" on animal ochdvior that have been w ritt<>J1 over Ill<' last I<.>W years. 1ndt>e, a fig"... thaI unde""brnalt"!' the actual inc",. ... bi>cau,<, the iuunt/ll substantially enlarged its p.lge siu in l'l'lll . \1o"-,<,,,('r, the quality of tho> work, d" well as it, qU.Jnlity, to,.ch.l.nf:"d. The a,-.,tage """"'I paper on ~nimal bet"",ior i. mo", "'Phi,Ii.;",tt-d ~nd inl<'''-'Sting than tho> ~ver ~ge paP"'" wriU,'n in It", 1'1805. SCi......lists ....~ lly do build ,m what olh~", haH' acrom- plisl-...d, whi.;h ""tl; lho.' b.lr hi~l><'r and higl><'r ~Mh y~a,. A. ,,-'S<'a,,-'he.. haw ""'" the chal\e.-l~,' of ~xp., nding our al"-"'dy 'ub.t.mtial ""',,,,ledg.' ,,j animal b<'ruvior, theyhaw gt"Ot"'al<'tI an ~mNrra"""""'t of riches for a texlb""k " 'ril<'1". As I han> ",viS<'bk'm holS se<1ll00especially aeun-. ~vt"J1 the numbo'.. of ..xcel- 1...,1 ......arrh "'P',rts c"'-'pl''tI wilh my desire to limil th., I""gtb of my boo k, D<->pite <'Iforts in th., p"'1 10 k,,,-,p th~ number of pa~,.. of texl und~, " mImi. my book had grown bigger over the y""rs. nu. lime Jhav~ m.u..g,>d 10",verse the lnend, which should make the book '. conn-nls mo", dig,'Stibl", although it d,,,.. m..an that "'''"Cgood ,tori,.. w......omitted. Read~ 1'!l "f my book can, ho"'~\'('r, ,'xplo", any b<'1""'ioral tOPKoj Iht-ir choice by turning 10 tho> origi- nallitetatu",. In this ""Hant onE" tool-the Web of Science (htlp: //www,w..oo/- 5Ciel\C<'.com)-il; lrt'm('lldo usly valuabl.. because il proviJ... such ~n ~~sy and quirk ",ay 10 Irack down lhe wurk of particular "-'SC'an:ht """",mrn''fld this ...arch me<:hani'm 100 sln>ngly. In addition 10 b , in~ng Ih~ boo k up 10 dat.. and """uclng ils I""gth ,,>rn,~ whal, Jhow" bc\>n abl.. to Inrorrx-"'dl~ ....,1'" illustration. ltiroul':h<-.ul tho> text, Ihanks 10 Ih~ willingn..," of mv publi.her, Sinau~r A''''lC;at...., to m~ k.. th.. mow f,om hlMk and ",hitt, to full rolor.This change not only add, 10Ih<' a,..- thelK val"" of tho.' book, bUIshuuld .1"" ht-lp ,....d...,. gr.,-;.p tl><' poinl of graphs xu P,e/ace more quic kly and easHy. The photograph. may also help .tudents why so many bclt",ioral biologi.l.low .lI.ldyinlll'l'al anilT4lls in ,..,al''1wiroror nts. The many dI;"""mple. 01tht' wonderfully inte""ting ........reh d,me on animal beh.vior will ft\dke the point th.t sci",,· tiflC logic offers a powerful means to gain an underst.wding of nalure, Acknowledgments The author of.ny texlbook depends on cooperalion from a surpri.ingly lal')\" number of pe",p l" . A, I have rewriU"" lhe bo,l<,k again, I haw, been very for- tun. le in having many generous colle. g ue. willing 10 give me th..ir lime, advice, pho togr aphs, and other form. of hdp, Mike Maxwell was especially helpful in providing informalion on m;c" ldlile analy" ",. All the cha pten< of th e boo k have incorporated chang uggested 10 me by Ihe following revi"we....: Alex IMsolo, Eliot Brenowilz, Ken Cat.nia, Robin Dunbar, Bruce Lyon, Jim Marden, Bob Montgom..rie. Randy Nd"m, Gabrielle Nevill. Don Owings, Steve '\Iowkki, Ko:m JU.e, .." G:n<' Robin""', Tom Seek>y, Nancy Segal, David ""'''''!neat, and leanne Zeh. Special thanks to Bob Montgomerie for read· ing tht' entire manu"':Tipt and for offering "x!remely helpful .uggestions about how 10do thin"" better. A host of other colleagu... have provided m" wit h f""rmission to use iIIu<-- tration. that originally apf""a""" in their papers. and some have given me the iIIuslrations !h<'m...i"..... I wanl to singk- out Bruce l yon for speciallhanl:rbphotog raph. ap!"'ar in many pta"" throughout the text as well a. on the cover of the boo k, I have ackno wledged all supplie... of pholographs in the !<>xl allhe appropriate figul'l'S.Acknowledgments 10the pu bli.hers who ha"e also g''Jl,,,ously granted f""nnissi,m to use their copyri l<;hted material .ppe.ron p.a~ located between the Bibliography .nd the Index. ~fy editor al Sin.""r A.sociates, Pete Farley, has done the hard and often unglamourous work of keeping the project mov ing ahead. I am wry grateful to him as well a. to all the other Sina uerian•. especially CheL.... Holabird, J"an Gemme, Chris Small, David Mcintyre, and Mara Sil"er, Norma Roche, who copyedl ted this edition as well as many othe.... ,..,ally know~ how to fix mis- takes. Readers of my boo k are luc ky thallh<'}' get to read lhe copyedited ver· sion rather than the original dr.ft, Although dII tog<'lher at lunch and sometim... on Friday aft"r- ....... iDrt-.. juot ~ _ t..,...toowl. doing iDrdoocada AdmitWly.1Iwamount cl ........ INI _ (OMUmi" Iwo ~ . dIOWb,& from tt... pildwr P""" pn"iOn cl tt... good old d.I~ 10. sJ-or two rurTmtly. WI tt-e \IIft o..-d by driIIoIo~) t..,.............lrloI"d IIw ........... INnk goo

Animal Behavior: An Evolutionary Approach

Similar Animal Behavior: An Evolutionary Approach