AN INTRODUCTION TO ANIMAL BEHAVIOUR Sixth Edition Wolvesexcitedlygreeteachotherasmembersofthepackcometogether;abumblebeeusesitslong tonguetoreachthenectaratthebaseofafoxgloveflower;aperegrinefalconstoopsathighspeedto strikeapigeonflyingbelow;youngcheetahsrestquietlytogether,veryclosetosleep. Nowinfullcolour,thisrevisedandupdatededitionofManningandDawkins’classictextprovides abeautifullywrittenintroductiontothefundamentalsofanimalbehaviour.Tinbergen’sfourquestions ofcausation,evolution,developmentandfunctionformthefundamentalframeworkofthetext, illustratedwithfascinatingexamplesofcomplexbehaviouralmechanisms.Theauthorsprovideaccounts ofalllevelsofbehaviourfromthenervecelltothatofthepopulation.ThestrengthsofAnIntroductionto AnimalBehaviourasatextbookincludeitsclearexplanationsandconcise,readabletext,andthe enthusiasmoftheauthorsfortheirsubject. AUBREY MANNING, OBE FRSE wasProfessorofNaturalHistoryattheUniversityofEdinburgh. Hismainresearchandteachinginterestswerebasedaroundthedevelopmentandevolutionofanimal behaviour.HecompletedhisDPhilunderNikoTinbergenandisnowEmeritusProfessor.Hereceived theZoologicalSocietyofLondonSilverMedalin2003forpublicunderstandingofscience. MARIAN STAMP DAWKINS isProfessorofAnimalBehaviourattheUniversityofOxfordwhereshe headstheAnimalBehaviourResearchGroup.Herresearchinterestsliearoundanimalperceptualworlds, withparticularemphasisonanimalsuffering.Shehasauthored,co-authoredandeditedeightbooks dealingwithsubjectsrangingfrombasicbehaviourtoanimalsufferingandconsciousness. AN INTRODUCTION TO ANIMAL BEHAVIOUR Sixth Edition Aubrey Manning UniversityofEdinburgh Marian Stamp Dawkins UniversityofOxford CAMBRIDGEUNIVERSITYPRESS Cambridge,NewYork,Melbourne,Madrid,CapeTown, Singapore,Sa˜oPaulo,Delhi,MexicoCity CambridgeUniversityPress TheEdinburghBuilding,CambridgeCB28RU,UK PublishedintheUnitedStatesofAmericabyCambridgeUniversityPress,NewYork www.cambridge.org Informationonthistitle:www.cambridge.org/9781107000162 #AubreyManningandMarianStampDawkins2012 Thispublicationisincopyright.Subjecttostatutoryexception andtotheprovisionsofrelevantcollectivelicensingagreements, noreproductionofanypartmaytakeplacewithout thewrittenpermissionofCambridgeUniversityPress. Firstpublished2012 PrintedintheUnitedKingdomattheUniversityPress,Cambridge AcataloguerecordforthispublicationisavailablefromtheBritishLibrary LibraryofCongressCataloging-in-PublicationData Manning,Aubrey. Anintroductiontoanimalbehaviour/AubreyManning,MarianStampDawkins.–6thed. p. cm. Includesbibliographicalreferencesandindex. ISBN978-1-107-00016-2(Hardback)–ISBN978-0-521-16514-3(Paperback) 1. Animalbehavior. I. Dawkins,MarianStamp. II. Title. QL751.M222012 591.5–dc23 2011027496 ISBN978-1-107-00016-2Hardback ISBN978-0-521-16514-3Paperback Additionalresourcesforthispublicationatwww.cambridge.org/9781107000162 CambridgeUniversityPresshasnoresponsibilityforthepersistenceor accuracyofURLsforexternalorthird-partyinternetwebsitesreferredto inthispublication,anddoesnotguaranteethatanycontentonsuch websitesis,orwillremain,accurateorappropriate. CONTENTS Preface pageix 1 Introduction 1 Questionsaboutanimalbehaviour 5 Theescapingcockroach 7 Thecourtshipofthesagegrouse 11 Unitsofthenervoussystem 16 Reflexesandmorecomplexbehaviour 20 Diversityandunityinthestudyofbehaviour 30 Summary 31 2 The development of behaviour 33 Younganimalsgrowup 34 Instinctandlearningintheirbiologicalsetting 40 Thecharacteristicsofinstinctandlearning 45 Geneticsandbehaviour 49 Developmentandchangestothenervoussystem 57 Hormonesandearlydevelopment 62 Earlyexperienceandthediversityofparentalbehaviour 68 Play 78 Imprinting 84 Birdsongdevelopment 94 Conclusions 106 Summary 106 3 Stimuli and communication 109 Whatstimuliareandhowtheyact 111 Diversesensorycapacities 114 v vi Contents Theproblemofpatternrecognition 126 Signstimuli(keyfeatures) 128 ‘Supernormal’stimuli 130 Neuroethologicalbasisofsignstimuli 135 Othersolutionsforpatternrecognition:generalizedfeaturedetection 139 Communication 144 Whatiscommunication? 145 Animalsignalsaseffectivestimuli 150 Honestyanddeceptioninanimalsignalling 159 Thehoneybeedance 164 Thecallsofvervetmonkeys 173 Summary 177 4 Decision-making and motivation 179 Decision-makingondifferenttimescales 182 Decision-makingand‘motivation’ 193 Measuringmotivation 193 Ismotivationspecificorgeneral? 199 Goalsasdecisionpoints 202 Homeostasisandnegativefeedback 205 Competitionbetweenmotivations 209 Inhibition/disinhibition 211 Decision-makingwithincompleteinformation:theroleofsignals 213 Conflictand‘abnormal’behaviour 217 Thephysiologyofdecision-making 220 Hormonesandsequencesofbehaviour 227 Conflictandphysiologicalstress 232 Decision-making,motivationandanimalwelfare 233 Conclusions 236 Summary 237 5 Learning and memory 239 Learningaspartofadaptation 239 Sensitizationandhabituation 242 Associativelearning 246 Specializedtypesoflearningability 257 Whatdoanimalsactuallylearn? 262 Aretherehigherformsoflearninginanimals? 264 Thecomparativestudyoflearning 268 Sociallearningandculture 272 Thenatureofanimalminds 276 Contents vii Thenatureofmemory 294 Summary 298 6 Evolution 301 Theadaptivenessofbehaviour 302 Genesandbehaviouralevolution 310 Kinselectionandinclusivefitness 316 Evolutionarilystablestrategies 328 Sexandsexualselection 336 Speciesisolationandspeciesselection 345 Tinbergen’sfourthquestion:thephylogenyofbehaviour 348 Summary 353 7 Social organization 355 Theindividualinthecrowd 355 Advantagesofgrouping 358 Diversesocialgroups 364 Eusociality:divisionintocastes 364 Territoryinthesocialorganizationofvertebrates 369 Matingsystemsandsocialorganization 373 Dominanceinsocialsystems 378 Diversemammaliansocialbehaviour 381 Primatesocialorganization 388 Summary 403 References 405 Figurecredits 435 Index 442