THE COGNITIVE ANIMAL This page intentionally left blank THE COGNITIVE ANIMAL Empirical and Theoretical Perspectives on Animal Cognition edited by Marc Beko¤, Colin Allen, and Gordon M. Burghardt ABradfordBook TheMITPress Cambridge,Massachusetts London,England (2002MassachusettsInstituteofTechnology Allrightsreserved.Nopartofthisbookmaybereproducedinanyformbyanyelectronicormechanicalmeans (includingphotocopying,recording,orinformationstorageandretrieval)withoutpermissioninwritingfromthe publisher. ThisbookwassetinTimesNewRomanon3B2byAscoTypesetters,HongKong.PrintedandboundintheUnited StatesofAmerica. LibraryofCongressCataloging-in-PublicationData Thecognitiveanimal:empiricalandtheoreticalperspectivesonanimalcognition/ editedbyMarcBeko¤,ColinAllen,andGordonM.Burghardt. p. cm. ‘‘ABradfordbook.’’ Includesbibliographicalreferences. ISBN0-262-02514-0(hc.:alk.paper)—ISBN0-262-52322-1(pbk.:alk.paper) 1.Cognitioninanimals. I.Beko¤,Marc. II.Allen,Colin. III.Burghardt,GordonM., 1941– QL785.C4852002 591.5013—dc21 2001057965 Contents Introduction ix 11 LearningandMemoryWithouta Contributors xvii Brain 77 JamesW.Grau I THE DIVERSITY OF 12 CognitiveModulationofSexual COGNITION 1 Behavior 89 MichaelDomjan 1 TheInnerLifeofEarthworms: Darwin’sArgumentandIts 13 CognitionandEmotioninConcert Implications 3 inHumanandNonhumanAnimals 97 EileenCrist RuudvandenBos,BartB.Houx, andBerryM.Spruijt 2 Crotalomorphism:AMetaphorfor UnderstandingAnthropomorphism 14 ConstructingAnimalCognition 105 byOmission 9 WilliamTimberlake Jesu´sRivasandGordonM. 15 Genetics,Plasticity,andthe Burghardt EvolutionofCognitiveProcesses 115 3 TheCognitiveDefender:How GordonM.Burghardt GroundSquirrelsAssessTheir 16 SpatialBehavior,FoodStoring,and Predators 19 theModularMind 123 DonaldH.Owings SaraJ.Shettleworth 4 JumpingSpiderTricksters:Deceit, 17 SpatialandSocialCognitionin Predation,andCognition 27 Corvids:AnEvolutionaryApproach 129 StimWilcoxandRobertJackson RussellP.BaldaandAlanC. 5 TheUngulateMind 35 Kamil JohnA.Byers 18 EnvironmentalComplexity,Signal 6 CanHoneyBeesCreateCognitive Detection,andtheEvolutionof Maps? 41 Cognition 135 JamesL.Gould PeterGodfrey-Smith 7 RavenConsciousness 47 19 CognitionasanIndependent BerndHeinrich Variable:VirtualEcology 143 AlanC.KamilandAlanB.Bond 8 AnimalMinds,HumanMinds 53 EricSaidel 20 SyntheticEthology:ANewToolfor InvestigatingAnimalCognition 151 9 ComparativeDevelopmental BruceMacLennan EvolutionaryPsychologyand CognitiveEthology:Contrastingbut 21 FromCognitioninAnimalsto CompatibleResearchPrograms 59 CognitioninSuperorganisms 157 SueTaylorParker CharlesE.Taylor 10 CognitiveEthologyattheEndof 22 ConsortTurnoversasDistributed Neuroscience 69 CognitioninOliveBaboons:A DaleJamieson SystemsApproachtoMind 163 DeborahForster Contents vi II CONCEPTS AND CATEGORIES 173 33 MeaningfulAcousticUnitsin NonhumanPrimateVocalBehavior 265 23 GeneralSigns 175 CoryT.MillerandAsifA. EdwardA.Wasserman Ghazanfar 24 TheCognitiveDolphin 183 34 ExploringtheCognitiveWorldof HerbertL.Roitblat theBottlenosedDolphin 275 LouisM.Herman 25 ChimpanzeeAiandHerSon Ayumu:AnEpisodeofEducationby 35 ChimpanzeeSigning:Darwinian Master-Apprenticeship 189 RealitiesandCartesianDelusions 285 TetsuroMatsuzawa RogerS.Fouts,MaryLeeA. Jensvold,andDeborahH.Fouts 26 TheEvolutionandOntogenyof OrdinalNumericalAbility 197 36 PrimateVocalandGestural ElizabethM.BrannonandHerbert Communication 293 S.Terrace MichaelTomaselloandKlaus Zuberbu¨hler 27 Domain-SpecificKnowledgein HumanChildrenandNonhuman 37 GesturalCommunicationinOlive Primates:ArtifactsandFoods 205 BaboonsandDomesticDogs 301 LaurieR.Santos,MarcD.Hauser, BarbaraSmuts andElizabethS.Spelke 38 AnimalVocalCommunication:Say 28 TheCognitiveSeaLion:Meaning What? 307 andMemoryintheLaboratoryand DrewRendallandMichaelJ. inNature 217 Owren RonaldJ.Schusterman,Colleen 39 CrackingtheCode:Communication ReichmuthKastak,andDavid andCognitioninBirds 315 Kastak ChristopherS.Evans 29 Same-Di¤erentConceptFormation inPigeons 229 IV SELF AND OTHER: THE RobertG.Cook EVOLUTION OF COGNITIVE COOPERATORS 323 30 CategorizationandConceptual BehaviorinNonhumanPrimates 239 40 TheMirrorTest 325 JacquesVauclair GordonG.Gallup,Jr.,JamesR. 31 CognitiveandCommunicative Anderson,andDanielJ.Shillito AbilitiesofGreyParrots 247 41 WhenTraditionalMethodologies IreneMaxinePepperberg Fail:CognitiveStudiesofGreat Apes 335 III COMMUNICATION, RobertW.ShumakerandKarylB. LANGUAGE, AND MEANING 255 Swartz 32 CognitionandCommunicationin 42 Kinesthetic-VisualMatching, PrairieDogs 257 Imitation,andSelf-Recognition 345 C.N.Slobodchiko¤ RobertW.Mitchell Contents vii 43 Darwin’sContinuumandthe 53 TheEvolutionofSocialPlay: BuildingBlocksofDeception 353 InterdisciplinaryAnalysesof Gu¨venGu¨zeldere,EddyNahmias, CognitiveProcesses 429 andRobertO.Deaner MarcBeko¤andColinAllen 44 IntegratingTwoEvolutionary 54 TheMoralsofAnimalMinds 437 ModelsfortheStudyofSocial LoriGruen Cognition 363 55 EyeGazeInformation-Processing BrianHareandRichard Theory:ACaseStudyinPrimate Wrangham CognitiveNeuroethology 443 45 FieldStudiesofSocialCognitionin BrianL.Keeley SpottedHyenas 371 56 TheEyes,theHand,andtheMind: KayE.HolekampandAnneL. BehavioralandNeurophysiological Engh AspectsofSocialCognition 451 46 TheStructureofSocialKnowledge VittorioGallese,PierFrancesco inMonkeys 379 Ferrari,EvelyneKohler,and RobertM.SeyfarthandDorothy LeonardoFogassi L.Cheney 57 VigilanceandPerceptionofSocial 47 FromtheFieldtotheLaboratory Stimuli:ViewsfromEthologyand andBackAgain:Cultureand SocialNeuroscience 463 ‘‘SocialMind’’inPrimates 385 AdrianTrevesandDiegoPizzagalli AndrewWhiten Afterword:WhatIsItLike? 471 48 EvolutionaryPsychologyand DonaldR.Gri‰n PrimateCognition 393 RichardW.Byrne Index 475 49 HowSmartDoesaHunterNeedto Be? 399 CraigB.Stanford 50 InsightfromCapuchinMonkey Studies:Ingredientsof,Recipesfor, andFlawsinCapuchins’Success 405 ElisabettaVisalberghi 51 ACognitiveApproachtotheStudy ofAnimalCooperation 413 LeeAlanDugatkinandMichaelS. Alfieri 52 KeepinginTouch:PlayFighting andSocialKnowledge 421 SergioM.Pellis This page intentionally left blank Introduction There are as many approaches to studying ani- andThe Expression of theEmotions in Man and mal cognition as there are definitions of cogni- Animals (1872). Consequently, both disciplines tionitself.Thisdiversityisreflectedintheessays are almost inextricably linked to the concept of that follow, to a degree that we believe is un- instinct. Darwin viewed instinct primarily in be- paralleled in any other volume that has been havioral terms and considered his ability to ex- produced onthissubject. This diversity is philo- plaininstinctthroughnaturalselectiontobeone sophical and methodological, with contributors ofthe mostcritical tests ofhistheories. Thus he demonstrating various degrees of acceptance or compared closely related species of bees to ex- disdain for terms such as ‘‘consciousness’’ and plain the evolution of hive building and closely various degrees of concern for the rigors of lab- related species of ants to explain the origins of oratory experimentation versus the validity of slave making. He also focused on domesti- naturalisticresearch.Thediversityisalsoappar- cated species to show how human intervention ent in the wide range of species to be found be- and selection could have formed variants. His tween these covers. Of course there are many prime examples here were di¤erent breeds of chapters on primates, and especially the great dogsandpigeons.Soon,zoologistssuchasC.O. apes, reflecting our human-centered interest in Whitman,E.C.Poulton,OskarHeinroth,Julian our closest relatives, but there are also chapters Huxley, and others began to exploit the com- that touch on cognition in animals as diverse as parative method to trace the evolution of even earthworms, antelopes, dogs, spiders, dolphins, the most complex social displays of animals. bees,fish,hyenas,snakes,sealions,prairiedogs, This work inspired the development of ethology virtual organisms, parrots, rats, ravens, and primarily through the research and promotional squirrels, to name but a few. We think we have e¤orts of the Nobel laureates Konrad Lorenz produced one of the most wide-ranging menag- andNikoTinbergen. eriesofscientificstudiesofanimalcognitionever Although when the Origin appeared in 1859 assembled. Yet it is humbling to realize that in Darwin deliberately discussed neither the evolu- terms of the diversity of life itself we cannot tion nor the behavior of human beings, he did, claimtohaveevenscratchedthesurface. neartheveryendofthebook,includethisintri- One of our objectives in bringing such a di- guingpassage: versecollection ofresearchstudiestogetheristo InthedistantfutureIseeopenfieldsforfarmoreim- show that whatever the ideological di¤erences, portantresearches.Psychologywillbebasedonanew behavioristicpsychologistsandcognitiveetholo- foundation,thatofthenecessaryacquirementofeach gists have a lot to learn from each other and mentalpowerandcapacitybygradation.Lightwillbe fromtheneurosciences.Yestherearedi¤erences thrownontheoriginofmanandhishistory.(Darwin ofopinionabouthowtopursuethestudyofan- 1859,p.488) imal cognition, but all sides stand to gain from listening carefullytotheconcerns ofothers.De- Note that this passage considers the psychologi- spite the di¤erences, there is in fact a great deal calaspectsofevolutionasfarmoreimportant,in ofsimilarityamongthedi¤erentapproaches,for the long run, than the morphological and taxo- theyhave,afterall,evolvedfromthesamestart- nomicissuestowhichtheOriginwaslargelyde- ing point in the theory of evolution by natural voted. Most biologists have ignored the clear selection. evidence, found in Darwin’s early notebooks, The roots of both comparative ethology and that while Darwin may have loved natural his- comparative psychology are found in the writ- tory, his ultimate agenda was to explain the ings of Charles Darwin, particularly in the Ori- origin of perhaps the strangest species of all, ginofSpecies(1859),TheDescentofMan(1871), ourselves. Thus, in his later writings Darwin
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