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392 Pages·2004·9.793 MB·English
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COMPARATIVE VERTEBRATE COGNITION: ARE PRIMATES SUPERIOR TO NON-PRIMATES? DEVELOPMENTS IN PRIMATOLOGY: PROGRESSAND PROSPECTS Series Editor: Russell H. Tuttle UniversityofChicago, Chicago, Illinois This peer-reviewed bookseries will meld the facts oforganic diversity with the continuity ofthe evolutionaryprocess. The volumesin thisserieswillexemplifythe diversityoftheo retical perspectives and methodological approaches currently employed by primatologists and physical anthropologists. Specific coverage includes: primate behavior in natural habi tats and captive settings: primate ecology and conservation; functional morphology and developmental biologyofprimates;primatesystematics;geneticandphenotypicdifferences among living primates;and paleoprimatology. ALLAPES GREATAND SMALL VOLUME 1: AFRICANAPES Edited byBirute M. F. Galdikas, Nancy Erickson Briggs, Lori K. Sheeran, Gary L. Shapiro and Jane Goodall THE GUENONS: DIVERSITYAND ADAPTATION IN AFRICAN MONKEYS Edited by Mary E. Glenn and Marina Cords ANIMALBODIES, HUMAN MINDS: APE, DOLPHIN,AND PARROTLANGUAGE SKILLS WilliamA. Hillixand Duane M. Rumbaugh COMPARATIVEVERTEBRATE COGNITION: ARE PRIMATES SUPERIORTO NON-PRIMATES? LesleyJ. Rogers and Gisela Kaplan COMPARATIVE VERTEBRATE COGNITION: ARE PRIMATES SUPERIOR TO NON-PRIMATES? Edited by Lesley J. Rogers Gisela Kaplan and Centre tor Neuroseienee and Animal Behaviour Sehool o[ Biologieal, Biomedieal and Moleeular Seienees University o[ New England Armidale, NSW 2351, Australia SPRINGER-SCIENCE+BUSINESS MEDIA, LLC Library ofCongress Cataloging-in-Publication Data Comparative vertebrate cognition: are primates superior to non-primates? / edited by Lesley ]. Rogers and Gisela Kaplan. p. cm. Includes bibliographical references and index. ISBN 0-306-47727-0 1. Cognition in animals. 2. Primates-Psychology. 3. Vertebrates-Psychology. I. Rogers, Lesley J. II. Kaplan, Gisela T. QL785.C537 2003 596.1513-dc22 2003060171 ISBN 978-1-4613-4717-0 ISBN 978-1-4419-8913-0 (eBook) DOI 10.1007/978-1-4419-8913-0 © 2004 Springer-Science+Business Media New York Originally publishcd by Kluwer Academic/Plenum Publishers, New York in 2004 Softcover reprint of the hardcover 1s t edition 2004 10 9 8 7 6 5 4 3 2 1 A C.I.P record for this book is available from the Library of Congress All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. PREFACE Primates have featured in humancultural history for alongtime. Adventurers, hunters, and travelers described them in their diaries. Even before Darwin, there was some recognition that they were different and fascinating, although not necessarilylikeable. Rennie, in his observation oforangutans in 1838 con demned orangutans as slovenly and useless creatures: "Their deportment is grave and melancholy, their disposition apathetic, their motions slow and heavy, and their habits so sluggish and lazy, that it is only the cravings of appetite, or the approach ofimminent danger, that can rouse them from their habitual lethargy, or force them to active exertion." (cit. in Yerkes, R. M., and Yerkes, A. W. (1929) The Great Apes. A Study of Anthropoid Life, Yale University Press, New Haven). Starting with Darwin, interest in primates became scientific and, tragically, led to the first major holocaust ofapes, who were killed by their thousands in short spans oftime to supply museums with specimens. Gradually zoos also captured them in increasing numbers for "exhibits." Because of their relatedness to humans, primates were then also regarded as good testsubjects in biomedical research and the beginningofthe trade and supply of monkeys commenced, taking them from the wild in uncontrolled and ever growing numbers for use in medical laborarories throughout the best part ofthe 20th century. Against this background, the efforts that were made in the 1960s and throughout the 1970s to say that primates and, in particular great apes, were "special" was amuch needed and radical departure from the poorpractice and derogatory attitudes that had gone before. Attitudes began to change along with the sudden rise to fame ofthe great apes, thanks to the extremely devot ed research work done by a number ofresearchers, many ofwhom are now household names. These changes resulted in questioning the ethics ofhowwe treated primates. Export and import of many primates is now banned in most countries; hence, taking them from the wild is often no longeranoption, no matter how justifiablethe purposemay be. Poachingis usuallypunishable bylaw(although v Preface VI notalwaysenforceable) and use ofprimates for frivolous researchpurposes has been curbed over the years. Overall, primates are now better protected in the wild and are generally treated betterin captivity than theywere in the 1950s. Such newappreciation can be attributed in no small measure to primatology's own hard work in disseminating the persistent take-home message that primates are special, deserving ofour interest, protection and moral support. As a result ofthese sustained efforts in the last 50 years or so, the image ofgreat apes gradually changed from that ofbrutes like King Kong (gorilla), the silly clown (chim panzee), or the slovenly imbecile (orangutan) to that ofcreatures worthy of appreciation or, less needed, we believe, ofsome kind ofnew "noble savage" perhaps even with amind ofits own. So what are we doing here in Primatology by publishing a book seemingly questioning the very essence on which we have finally achieved some change in moral status for primates? There are two points to be made in reply. The appalling past treatment ofprimates and their depletion in the wild had to be fought bypolitical means and discourses in the political realms ofhumansoci eties. The claim that primates are special was an important political point of argument; and so was the evolving concept ofgreat apes as flagship species. This book,voicingthe opinionsofsomeleadingprimatologists,ethologists, psychobiologists,neuroscientistsandanthropologists,isnotspeakingfrom the standpoint ofapolitical engagement with primates but ofa scientific engage ment with primates in relation to all other species. The extremely lively and seminal primatological debates ofthe 1970s, especially on the cognitive skills ofapes, have carried us almost through to this day. Thirtyyears later, itis time to take stock and revise what we know, casting the netwider than the primate orderto see what we have achieved theoreticallyandwhatis sustainable scien tifically. Such stock taking exercises help us to adjust our research directions, reflect on our practices, and perhaps also give rise to new and different questions. Ofteninadvertently, butsometimesdeliberately,we make statementsimply ingsuperiorityofthe behaviororcognitive abilities ofapes compared to mon keys, or primates compared to nonprimates. Yet recent evidence shows that at least some ofthe characteristics that we thought made primates unique and superior are alsoseenin some species ofnon-primates. Thus we are beginning to become aware ofthe risks ofmoving in one intellectual milieu and choos ing to work on aspecies instead ofchoosing aproblem and then testing it on Preface vii a range of species. With this recognition has come another motivation for editing this book. Each contributor has addressed the question "Are the cog nitive abilities ofprimates simplydifferent from other mammals or are they an evolutionary advance on them?" or, more generally, "Are primates special?" Eachcontributorhas approached the questionfrom adifferentperspective and considered arange ofdifferent species. We have covered in this book research findings about awide span ofvertebrate species, all relevant to understanding primates. Of course, we recognize that there are differences in cognition between all species, including the various primates, but we have examined whether there is any substance behind the often stated or hinted assumption that primates are some sort ofimproved cognitive design and that, amongst primates, the apes are afurther improvement that foregrounded the evolution ofthe human brainandmind. We have approached this topicfully awareofthe difficulties in defining what we mean by higher cognition, or "intelligence," butbyourvariouswaysofdealingwith this uncertaintyaswe compareprimate and nonprimate species we believe we will lead to achange in thinking. The aim ofthe book is to explore afresh the long-standing interest, and emphasis on, the "special" capacities ofprimates. Some ofthe recent discov eries of the higher cognitive abilities of other mammals and also birds challenge theconceptthatprimates are special andeventhe view that the cog nitive abilityofapes is more advanced than that ofnon-primate mammals and birds. It is therefore timely to ask whether primates are, in fact, superior to non-primates, and to do so from a broad range ofperspectives. Itwill become apparent to the reader that researchers are only at the beginning ofthe search to find out whether primates are special and, ofcourse, by "special" is meant notjustdifferent,whichapplies to all biologicalcategories, but "better" in the ephemeral sense of being more like us and being cognitively superior to all other species. Some of the contributors conclude that all primates (or some primate species) do, indeed, have abilities thatseparate them from othermammals and that these abilities are clearlyspecial. Others argue thatprimates as agroup are no more special than arange ofother species, or they are special only because they are seen in relation to human primates. Our aim has not been to down play studies on primates, or to simply criticize them, in comparison to studies on nonprimates, but to present the results ofsimilar studies on primates and nonprimates to see whether any or many ofthe known cognitive abilities of primates are shared by some ofthe nonprimate taxa. Preface V1ll Divided into five sections this book deals with topics about higher cogni tion and howitis manifestedindifferent species, and also considers aspects of brain structure that might be associated with complex behavior. One important theme of this book is, ofcourse, the evolution ofhigher cognition and another is the applicability ofdifferent testing methods to dif ferent species. We would fall short if we attempted here to summarize the breadth ofareas covered by the chapters, since eachone is acomplete essayin itself. We hope they will be read to enlighten scientifically, rather than in any sense to lessenourrespectfor primatespolitically. At the veryleast, orperhaps the most, we hope that this volume will stimulate comparative research on behavior and so ensure that future dividing lines between species are erected onlywhen theirfoundations are in science and not the resultofeithercultural distortions or political gain. Wewouldliketoexpressourgratitudeto RussellTuttle,editoroftheseries, and Andrea Macaluso, editor with Kluwer, whose invitation to us to proceed with this book gave us the opportunity to explore our ideas further, and to thank Richard Andrew, Dario Maestripieri, and Michael Beran for their valuable comments on an earlier draft ofthe manuscript. We thank Leanne Stewartfor assistance in preparingthe index. We are mostgratefulto allofthe contributors for adding so many ideas and giving us so many enjoyable pages to read. Lesley J. Rogers and Gisela Kaplan April, 2003 CONTENTS Contributors xv PART I: COMPLEX COGNITION 1. Comparing the Complex Cognition of Birds and Primates 3 Nathan]. Emeryand Nicola S. Clayton Introduction 3 Why Might Primates be Superior to Non-Primates? 4 Comparing Birds and Primates 5 Primates have a Neocortex Larger than Predicted for their Body Size 5 Primates have an Expanded Prefrontal Cortex 8 Primates Demonstrate Social Learning and Imitation 9 Primates Understand Others' Mental States 14 Primates Display Insight, Innovation, and they Construct and Use Tools 23 Insight and Innovation 23 Manufacture and Use ofTools 25 Primates Utilize Symbolic and Referential Communication 27 Primates Demonstrate Elements ofMental Time Travel 29 The Retrospective Component-Episodic Memory 30 Do Animals have Episodic-like Memory? 31 Episodic-like Memory in Scrub-Jays 32 The Prospective Component-Future Planning 33 Is there any Evidence ofFuture Planning in Animals? 35 Food Caching by Scrub-Jays: A Candidate for Future Planning inAnimals? 36 The Perils ofPrimatocentrism and "Scala Naturae" 36 Uses and Abuses ofthe Ecological/EthologicalApproach to Cognition 38 IX x Contents Species Differences in Ecology and Cognition 39 Ethologicaily Relevant Stimuli are Difficult to Control 40 HowFar can the Natural Behavior of an Animal be Translated to the Laboratory? 41 The Great Divide: Awareness of"Self' 41 Is there a Case for Convergent Cognitive Evolution and Divergent Neurological Evolution? 45 Acknowledgments 46 References 46 2. Visual Cognition and Representation in Birds and Primates 57 Giot;gio Vallortigara Introduction 57 Integration and Interpolation ofVisual Information in the Spatial Domain 58 Integration andInterpolation ofVisual Information in the Temporal Domain 69 Representing Objects 72 Objects inSpace: Use ofGeometric and Nongeometric Information 77 Conclusions 84 Acknowledgments 85 References 85 PART II: SOCIAL LEARNING 3. Socially Mediated Learning among Monkeys and Apes: Some Comparative Perspectives 97 Hilary O. BoxandAnneE. Russon Introduction 97 Socially Mediated Learning 98 Imitation 106 Imitation in Monkeys 106 Imitation in Great Apes UO Primate Imitation in Broader Perspective U5 Culture U8 Behavioral Traditions among Monkeys U9 Behavioral Traditions among the GreatApes 121

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