Table Of Content

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 1JIrAd<Ipt4t...."" ApyI_1I J5 I'.'lryDoOnly ,"",In: SINg ~ 39 N/ltrmlJ S<-I«tiorl.Pl4 o..I«ts 42 Proximal.. and L%m.tle c.~ Are Complementory 45 3 The DevelopmenrofBehavior:A Focus on Heredity 50 TIw Go.onetic5 of Beh.....io! 52 Gmt1i(" Dilfrrrnas Imd H......." Bt/Ion"", 55 Goorr.1ic Diffrrmcn ",,<I IQ f),~ 57 H_ Mmy Cmdic DIjfrrrncn A" NtrdftI to Pralwcr • &lwt.....aJ CJiffrrma~ S9 1l\I' h olulion 01Bt>h.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.<Jfipllwll fM 131 TIlt Pron""," &s;" of 5ti","II~ fillm"g J31 $ttttIoJ... FiI/niJIg '" A"",1ory RlaJoton 132 5Ii","11I5 fil"""g i" titt' Tctil.- M<oit JJ4 A<Upti\~ Prlninwte"~ of 80"'••1\_ 137 AMptiPrMtdlm...... of,...."uitwl 141 All OIji1ctory Map 145 6 I The Organization ofBehavior; Neurons and Hormones 1S4 How N..ural Command C..ntM'! OrganiZt' Ikha\';ur 156 CW MtduIllisms.lId 8rlrtlIOt>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""":<l'tIl EPOiw,ioll 190 The Di....rsity of Antipl't'darn. Ada ptations 192 ~ Va/wt ".f Body "Dtroralioll.'· 194 TItt V.I....ofW/lTIi'lig ~ 197 H<JW To51",.. Pun..". 200 TItt V.llIt ".f Vrgi"'lIO' 202 Tht Dillllioll E/ftclalld lilt StIfi..oJr Hmi 1Q.1 fightlllg Back 207 A Tactic of Lasl RI:S<,.,? 209 8 The Evolution ofFeeding Behavior 214 Incolhng Food 216 Tht 0rig11UofPr-ty-l..oal'"g MtdMfoisllts 117 Cortt,,,, Htlpfrom e-,..."'" 119 Tht History of~ &r DrlIIIaS 111 Tht A40tptin' Va.... afHonty &r £l,m,;r.o 225 Tht I~ U111" HypotltniJ 117 L«/JI'"g Prry I1y Dtmt 219 Sel«ting " 'hal lOEar 231 H_ '" Cltoost"" Opf.......a.r... 23J H_ '" Cltoost"" Opti-t MIIUd lJ4 Cnti<"is>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 Dr<prningfrom Onr P/lIa '" Anotlorr m M,gration 259 T1u' Cosl. ofMigralion 260 TIlt 1k'Nfit5 uf Migralitm 262 MigNfi<ln iIO .. QmdiliolW TiKti.:" 2U TemtorWity 266 TrrTiiorilllity ..ptJ c.lorits 269 TrrTilorUzl Puuks 271 Why Do Tf'Trill1'Y Ho/dm AI..."", Alw"y' Win? 272 10 The Evolution ofCommunication 280 The Origin. and Adaptive Value l>f a Sign<ll 282 C~...~lIIlit.. SrlKfitm 4P1d Mllllirk awngt5 283 TIlt AJoortiR' VoIlwuj hit o...ng<'< 284 Cllnmf AJoortItY V.w 286 11>0' History of .. Sign.al-Rf'mving M..dwusm 2'l7 n", History of1n"d Wing. 289 s"n.<lry hpl<litltiun of Signal Receiv,," by SigNlet's 292 x-y Ptrft',Joa'f ~ ~ flw £llCII~"'" of.. SigfwI 293 n", Panda P"n,ipk 297 The Ad"pta~tAppl'OlCh to CommWlK..tion ~m Yo1ry Oct &by Bmls&x SoN<tislJjIfor Food~ JOO Ukg'/I_ I~ Rt'l:tn",.s 30S Adapli,,,, Sig>1al Rt'Ui ,~n)( J07 Ricrivtrs Mily RI'1j~il'tH"IUS/ Sigrwls JOIj Why ~ lktJ'Iio>r O<nor~ JJI 11 The Evolution ofReproductive Behavior 316 "Jh!o holution CIt Dillt iE ....fl if! 5el< R'*'" 3111 Tnh"g IIw£llCIIIII'""""Y 11II't>fyujSn 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"-<tntial for Eusociality to [ 001,,,, 449 TIlt Ecology of[usocwlity 451 15 The Evolution ofHuman Behavior 456 1be Adaptalinnis! App roach 10 HUllliIn Beh.,"ior 4';8 TI,e SociobWlogyCo"t""",rsy 459 [1'lJlulum and the Diversity ofHuman Cult,,,,,, 462 Sociobiology versus Arbitrary Culture Theory 464 Adaptive Mating Oed sions 466 Adapli"" Mate Choice by Women 466 Glossary 489 Bibliography 495 lIIustrarion Credits 529 Index 53' Adapti!'" Mate Choice byMe" 471 Conflicl btfw«tl the Sexes 474 Coerci,,,,Sex 477 Adaptive Parental Care 4IlO Hel,'ing Chl/d,..n Marry 481 Preface A n"th~r f"ur y~" rs or so have passed ' in<:e the f'r<'viou" <.>dition 01 Animal Beha,,;")]" dppe.>n'd. Purin!\ thi, time, much thai L, d"Preso;ing hils occurn.'<l on II><' polilk<l1 and """;ronm.",,,,1 fronts_L..1 us IlOI dwell on the;e ""<'I1l>;, 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<I, the rMeat which impurt.,nt discowries about behaVK"-aTt' bt-inKmad" has st('..dily inc",.sed thn"'K"''''I Irn, Iit.,'lime of my textbook. Just pid ' up a n'O.",t ropy of any ;0"",,,1 in the dis<.~pli"" and compare the " ....arch reports tlw", with tlu _ in the ""'" journal 15 or 2Q years p,,,v;ou,ly_Thu•• in 1982. t.....journal A"i"",r Bmwwur publisht-d 1264f"' ges oI ...ports; in 20ClJ the t,,"'1 was 2180 raK'''>, 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<'<! my bo"k ag~in, my m~in difficully has N"-'" whalto I"",v~ "u!. nol who,t 10 put in. Although there ha ,'~ alway. M 'fl m~ny mo.... ~ood pa!"," than spaC<'a"ailablc fur my .ummari.... Ihis time anlUnd lho.· pn>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:h<o'" and 10folk'w the trail of p"p.,rs that rit,' a "-""'arch "'f'Ort of inu,resl. I ram">t """",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<Inl<;'" thaI I have made '" !hi,.."""th edition of my bo,l<,k ha,'e all bteoen ft\dde with one primary goa l in mind; 10hel p my reade......... how ........ rche... have been able 10 reach .., tisfaclory =Iu.ion. aboul how and why animal. do tht' things theydo. I hope thaI it>;"""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<s; his sup>: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 dI<Ipter " ·vi,,,,'e..., photographe...., editorial.talf, and presidenl. of lhe United Slates often change from ed ilion to edition. some Ihings slay lhe "" me, which p"wid.... a certain reassuring stability t" my life. My wife Sue continues 10 cope with my many idiosyncra.i..., .Iillli.tening thoughtfully whenever I vigorously del\OUJ\('(-' the likes of, say, Henry Kissing..r or Antonin Scalia, sliUwilling 10live in a cramped ~amp',rvan for month. at a lime w hom we are in W...tern Australia for .nother round of bee research. My younger son !\;ick is .till in town rr..mpe, Arizona), and he takes time olf from help- ing lho6<' acruse<! of driving under the influence 10help u. eat dim"" and play ping-pong, A. in the pa.t, he lets me win onee in a while so that I can retain som.. small measure of S<-'11.... t<'<'m, thaI most important of mod..", com- modities, My older son loe is dose .'nough (Albuquerque, !\lew Mexico) thaI he ra n join uS 00 occasion, helping me ma in lain the iUusion of being .ur- rounded by family, In addition, I .m happy to report that none of my friends at Ariwna SI.lte (among lhem OO"e Brown, Sl,'v.. Carroll, Jim Colli,,", Stuart Fi.her. Dave Pearson, and Ron RUlowski) has yet go"e IIIhis reward, which means tha I we can aU g<>I 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 \I<lf"O undford......... ....... (Ihr nWJWltlUO dN<h-,",,~ cl grttinp; oldo.-f MIdIhr trout>Ift o..-d by dri<rti,·.. <:<>IIoIo~) t..,.............lrloI"d IIw ........... INnk goo<tr-. TCI my f...,- i1y .and fnend,. INro. y"". , ..r\ ". . ' . , .,, ,, • ;. -, r -:: ~ • :.... : : - , ~ : . J ,,' .... ... t •• :• . J .. " . . • • '- ... ". • , ~ • . - • .. , ... (""Tk. Omwjn~ , rudy in Downtlou'" -'MQt-••!optd 'M r/IMfy of MlIurion by ""'u,,,i lfIIoailln ,n. faunda'ion fDItn. _m 'Mtyof anjmaJ~, Pf><J'''9raph by Ma,l; Mofftrr An Evolutionary Approach to Animal Behavior or hundreds of thousands of years, humans observed animals because their lives depended on a knowledge of animal behavior. Even today, the sub- ject still has great practical significance. Information on the reprod uctive behavior of insect pests, for example, may ultimately lead 10 their control, while knowledge of the migratory routes of an endangered whale or shorebird may enable conservanonlsts to design adequate reserves to save the animal from extinction. Moreover, an understanding of the evolutionary basis of our own behavior might help us identify why we so often damage our environmen ts, per- haps enabling us to reduce ou r destructive tendencies (1251). But even if the only beneficiaries of studies of ani- mal behavior were the persons who conducted the research, 1suspect that work in this field would continue. Learning how and why animals behave is an intrinsically fascina ting business. Perhaps you can imagine what it would be like to be the first person to discover that male damselflies actually use their penis as a scrub brush to remove the sperm of rival males from their mates [11 771; or maybe you can put yourself in the shoes of the person who first showed that female Seychelles warblers could control

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Animal Behavior: An Evolutionary Approach PDF

505 Pages·2001·65.12 MB·English
by  John Alcock
#animal behavior #animal behaviour
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Preview Animal Behavior: An Evolutionary Approach

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 1JIrAd<Ipt4t...."" ApyI_1I J5 I'.'lryDoOnly ,"",In: SINg ~ 39 N/ltrmlJ S<-I«tiorl.Pl4 o..I«ts 42 Proximal.. and L%m.tle c.~ Are Complementory 45 3 The DevelopmenrofBehavior:A Focus on Heredity 50 TIw Go.onetic5 of Beh.....io! 52 Gmt1i(" Dilfrrrnas Imd H......." Bt/Ion"", 55 Goorr.1ic Diffrrmcn ",,<I IQ f),~ 57 H_ Mmy Cmdic DIjfrrrncn A" NtrdftI to Pralwcr • &lwt.....aJ CJiffrrma~ S9 1l\I' h olulion 01Bt>h.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.<Jfipllwll fM 131 TIlt Pron""," &s;" of 5ti","II~ fillm"g J31 $ttttIoJ... FiI/niJIg '" A"",1ory RlaJoton 132 5Ii","11I5 fil"""g i" titt' Tctil.- M<oit JJ4 A<Upti\~ Prlninwte"~ of 80"'••1\_ 137 AMptiPrMtdlm...... of,...."uitwl 141 All OIji1ctory Map 145 6 I The Organization ofBehavior; Neurons and Hormones 1S4 How N..ural Command C..ntM'! OrganiZt' Ikha\';ur 156 CW MtduIllisms.lId 8rlrtlIOt>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""":<l'tIl EPOiw,ioll 190 The Di....rsity of Antipl't'darn. Ada ptations 192 ~ Va/wt ".f Body "Dtroralioll.'· 194 TItt V.I....ofW/lTIi'lig ~ 197 H<JW To51",.. Pun..". 200 TItt V.llIt ".f Vrgi"'lIO' 202 Tht Dillllioll E/ftclalld lilt StIfi..oJr Hmi 1Q.1 fightlllg Back 207 A Tactic of Lasl RI:S<,.,? 209 8 The Evolution ofFeeding Behavior 214 Incolhng Food 216 Tht 0rig11UofPr-ty-l..oal'"g MtdMfoisllts 117 Cortt,,,, Htlpfrom e-,..."'" 119 Tht History of~ &r DrlIIIaS 111 Tht A40tptin' Va.... afHonty &r £l,m,;r.o 225 Tht I~ U111" HypotltniJ 117 L«/JI'"g Prry I1y Dtmt 219 Sel«ting " 'hal lOEar 231 H_ '" Cltoost"" Opf.......a.r... 23J H_ '" Cltoost"" Opti-t MIIUd lJ4 Cnti<"is>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 Dr<prningfrom Onr P/lIa '" Anotlorr m M,gration 259 T1u' Cosl. ofMigralion 260 TIlt 1k'Nfit5 uf Migralitm 262 MigNfi<ln iIO .. QmdiliolW TiKti.:" 2U TemtorWity 266 TrrTiiorilllity ..ptJ c.lorits 269 TrrTilorUzl Puuks 271 Why Do Tf'Trill1'Y Ho/dm AI..."", Alw"y' Win? 272 10 The Evolution ofCommunication 280 The Origin. and Adaptive Value l>f a Sign<ll 282 C~...~lIIlit.. SrlKfitm 4P1d Mllllirk awngt5 283 TIlt AJoortiR' VoIlwuj hit o...ng<'< 284 Cllnmf AJoortItY V.w 286 11>0' History of .. Sign.al-Rf'mving M..dwusm 2'l7 n", History of1n"d Wing. 289 s"n.<lry hpl<litltiun of Signal Receiv,," by SigNlet's 292 x-y Ptrft',Joa'f ~ ~ flw £llCII~"'" of.. SigfwI 293 n", Panda P"n,ipk 297 The Ad"pta~tAppl'OlCh to CommWlK..tion ~m Yo1ry Oct &by Bmls&x SoN<tislJjIfor Food~ JOO Ukg'/I_ I~ Rt'l:tn",.s 30S Adapli,,,, Sig>1al Rt'Ui ,~n)( J07 Ricrivtrs Mily RI'1j~il'tH"IUS/ Sigrwls JOIj Why ~ lktJ'Iio>r O<nor~ JJI 11 The Evolution ofReproductive Behavior 316 "Jh!o holution CIt Dillt iE ....fl if! 5el< R'*'" 3111 Tnh"g IIw£llCIIIII'""""Y 11II't>fyujSn 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"-<tntial for Eusociality to [ 001,,,, 449 TIlt Ecology of[usocwlity 451 15 The Evolution ofHuman Behavior 456 1be Adaptalinnis! App roach 10 HUllliIn Beh.,"ior 4';8 TI,e SociobWlogyCo"t""",rsy 459 [1'lJlulum and the Diversity ofHuman Cult,,,,,, 462 Sociobiology versus Arbitrary Culture Theory 464 Adaptive Mating Oed sions 466 Adapli"" Mate Choice by Women 466 Glossary 489 Bibliography 495 lIIustrarion Credits 529 Index 53' Adapti!'" Mate Choice byMe" 471 Conflicl btfw«tl the Sexes 474 Coerci,,,,Sex 477 Adaptive Parental Care 4IlO Hel,'ing Chl/d,..n Marry 481 Preface A n"th~r f"ur y~" rs or so have passed ' in<:e the f'r<'viou" <.>dition 01 Animal Beha,,;")]" dppe.>n'd. Purin!\ thi, time, much thai L, d"Preso;ing hils occurn.'<l on II><' polilk<l1 and """;ronm.",,,,1 fronts_L..1 us IlOI dwell on the;e ""<'I1l>;, 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<I, the rMeat which impurt.,nt discowries about behaVK"-aTt' bt-inKmad" has st('..dily inc",.sed thn"'K"''''I Irn, Iit.,'lime of my textbook. Just pid ' up a n'O.",t ropy of any ;0"",,,1 in the dis<.~pli"" and compare the " ....arch reports tlw", with tlu _ in the ""'" journal 15 or 2Q years p,,,v;ou,ly_Thu•• in 1982. t.....journal A"i"",r Bmwwur publisht-d 1264f"' ges oI ...ports; in 20ClJ the t,,"'1 was 2180 raK'''>, 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<'<! my bo"k ag~in, my m~in difficully has N"-'" whalto I"",v~ "u!. nol who,t 10 put in. Although there ha ,'~ alway. M 'fl m~ny mo.... ~ood pa!"," than spaC<'a"ailablc fur my .ummari.... Ihis time anlUnd lho.· pn>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:h<o'" and 10folk'w the trail of p"p.,rs that rit,' a "-""'arch "'f'Ort of inu,resl. I ram">t """",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<Inl<;'" thaI I have made '" !hi,.."""th edition of my bo,l<,k ha,'e all bteoen ft\dde with one primary goa l in mind; 10hel p my reade......... how ........ rche... have been able 10 reach .., tisfaclory =Iu.ion. aboul how and why animal. do tht' things theydo. I hope thaI it>;"""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<s; his sup>: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 dI<Ipter " ·vi,,,,'e..., photographe...., editorial.talf, and presidenl. of lhe United Slates often change from ed ilion to edition. some Ihings slay lhe "" me, which p"wid.... a certain reassuring stability t" my life. My wife Sue continues 10 cope with my many idiosyncra.i..., .Iillli.tening thoughtfully whenever I vigorously del\OUJ\('(-' the likes of, say, Henry Kissing..r or Antonin Scalia, sliUwilling 10live in a cramped ~amp',rvan for month. at a lime w hom we are in W...tern Australia for .nother round of bee research. My younger son !\;ick is .till in town rr..mpe, Arizona), and he takes time olf from help- ing lho6<' acruse<! of driving under the influence 10help u. eat dim"" and play ping-pong, A. in the pa.t, he lets me win onee in a while so that I can retain som.. small measure of S<-'11.... t<'<'m, thaI most important of mod..", com- modities, My older son loe is dose .'nough (Albuquerque, !\lew Mexico) thaI he ra n join uS 00 occasion, helping me ma in lain the iUusion of being .ur- rounded by family, In addition, I .m happy to report that none of my friends at Ariwna SI.lte (among lhem OO"e Brown, Sl,'v.. Carroll, Jim Colli,,", Stuart Fi.her. Dave Pearson, and Ron RUlowski) has yet go"e IIIhis reward, which means tha I we can aU g<>I 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 \I<lf"O undford......... ....... (Ihr nWJWltlUO dN<h-,",,~ cl grttinp; oldo.-f MIdIhr trout>Ift o..-d by dri<rti,·.. <:<>IIoIo~) t..,.............lrloI"d IIw ........... INnk goo<tr-. TCI my f...,- i1y .and fnend,. INro. y"". , ..r\ ". . ' . , .,, ,, • ;. -, r -:: ~ • :.... : : - , ~ : . J ,,' .... ... t •• :• . J .. " . . • • '- ... ". • , ~ • . - • .. , ... (""Tk. Omwjn~ , rudy in Downtlou'" -'MQt-••!optd 'M r/IMfy of MlIurion by ""'u,,,i lfIIoailln ,n. faunda'ion fDItn. _m 'Mtyof anjmaJ~, Pf><J'''9raph by Ma,l; Mofftrr An Evolutionary Approach to Animal Behavior or hundreds of thousands of years, humans observed animals because their lives depended on a knowledge of animal behavior. Even today, the sub- ject still has great practical significance. Information on the reprod uctive behavior of insect pests, for example, may ultimately lead 10 their control, while knowledge of the migratory routes of an endangered whale or shorebird may enable conservanonlsts to design adequate reserves to save the animal from extinction. Moreover, an understanding of the evolutionary basis of our own behavior might help us identify why we so often damage our environmen ts, per- haps enabling us to reduce ou r destructive tendencies (1251). But even if the only beneficiaries of studies of ani- mal behavior were the persons who conducted the research, 1suspect that work in this field would continue. Learning how and why animals behave is an intrinsically fascina ting business. Perhaps you can imagine what it would be like to be the first person to discover that male damselflies actually use their penis as a scrub brush to remove the sperm of rival males from their mates [11 771; or maybe you can put yourself in the shoes of the person who first showed that female Seychelles warblers could control

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