Economics in Nature Since the development of game theory, the analysis of animal behaviour using the theories of economics has become a growing field of biological research in which models of games and markets play an important role. Studies of sexual selection, interspecific mutualism and intraspecific cooperation show that individuals exchange commodities to their mutual benefit; the exchange values of commodi- ties are a source of conflict, and behavioural mechanisms such as partner choice and contest between competitors determine the composition of trading pairs or groups. These ‘biological markets’ can be examined to gain a better understand- ing of the underlying principles of evolutionary ecology. In this volume scientists from different disciplines combine insights from economics, evolutionary biology and the social sciences to look at comparative aspects of economic behaviour in humans and other animals. Aimed primarily at evolutionary biologists and anthropologists, it will also appeal to psychologists and economists interested in an evolutionary approach. Ronald Noë is Professor of Ethology at the Université Louis-Pasteur at Strasbourg, France and holds a post at the Max-Planck-Institut, Seewiesen, Germany. His main research interest is the behavioural ecology of primates and in 1991 he founded the Taï Monkey Project in the Taï National Park, Ivory Coast. He has been editor of the journal Ethology since 1997. Jan A. R. A. M. van Hooff is Professor of Behavioural Biology at the University of Utrecht. His research focuses on the social behaviour and organisation of pri- mates and eusocial insects. He is a member of the Royal Netherlands Academy of Arts and Sciences and the Scientific Council of the Jean-Marie Delwart Foundation, Belgium. Peter Hammerstein is Professor in Organismic Evolution at the Theoretical Biology Laboratory of Humboldt University, Berlin. Besides being a biologist he has a background in game theory and mathematical economics. His main research interest is in the evolutionary analysis of conflict and cooperation at the level of genes and organisms. Economics in Nature Social Dilemmas, Mate Choice and Biological Markets Edited by Ronald Noë, Jan A. R. A. M. van Hooff and Peter Hammerstein CAMBRIDGE UNIVERSITY PRESS Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo Cambridge University Press The Edinburgh Building, Cambridge CB2 2RU, UK Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521650144 © Cambridge University Press 2001 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2001 This digitally printed first paperback version 2005 A catalogue record for this publication is available from the British Library Library of Congress Cataloguing in Publication data Economics in nature: Social Dilemmas, Mate Choice and Biological Markets / edited by Ronald Noë, Jan A. R. A. M. van Hooff, and Peter Hammerstein. p. cm. Includes bibliographical references and index. ISBN 0 521 65014 3 (hardback) 1. Social behaviour in animals – Economic aspects. 2. Human behavior – Economic aspects. 3. Psychology, Comparative. I. Noë, Ronald, 1951– . II. Hooff, J. A. R. A. M. van (Johan Antoon Reinier Alex Maria), 1936– . III. Hammerstein, Peter, 1949– . QL775.E27 2001 591.56–dc21 00-065154 ISBN-13 978-0-521-65014-4 hardback ISBN-10 0-521-65014-3 hardback ISBN-13 978-0-521-00399-5 paperback ISBN-10 0-521-00399-7 paperback Contents List of contributors page vii Preface ix Acknowledgments xi 1 Games and markets: economic behaviour in humans and other animals 1 Peter Hammerstein Part I Economic behaviour in social networks 2 Social dilemmas and human behaviour 23 Elinor Ostrom 3 Cooperation and collective action in animal behaviour 42 Charles L. Nunn and Rebecca J. Lewis 4 Conflict, reconciliation and negotiation in non-human primates: the value of long-term relationships 67 Jan A. R. A. M. van Hooff Part II Biological markets 5 Biological markets: partner choice as the driving force behind the evolution of mutualisms 93 Ronald Noë 6 The utility of grooming in baboon troops 119 Louise Barrett and S. Peter Henzi 7 The cleaner fish market 146 Redouan Bshary v vi contents 8 Modelling interspecific mutualisms as biological markets 173 Jason D. Hoeksema and Mark W. Schwartz Part III Mating markets 9 Human mate choice strategies 187 Boguslaw Paw-lowski and Robin I. M. Dunbar 10 How does mate choice contribute to exaggeration and diversity in sexual characters? 203 Andrew Pomiankowski and Yoh Iwasa 11 Information about sperm competition and the economics of sperm allocation 221 Geoffrey A. Parker and Mike A. Ball 12 The economics of male mating strategies 245 Robin I. M. Dunbar Index 270 Contributors Mike A. Ball Department of Mathematical Sciences, University of Liverpool, Liverpool L69 3BX, UK Louise Barrett School of Biological Sciences, University of Liverpool, Liverpool L69 3BX, UK Redouan Bshary Max-Planck-Institut für Verhaltensphysiologie, Seewiesen, Postfach 1564, 82305 Starnberg, Germany Robin I. M. Dunbar ESRC Research Centre in Economic Learning and Social Evolution, School of Biological Sciences, Nicholson Building, University of Liverpool, Liverpool L69 3BX, UK Peter Hammerstein Innovationskolleg Theoretische Biologie, Humboldt-Universität zu Berlin, Invalidenstr. 43, D-10115 Berlin, Germany S. Peter Henzi Behavioural Ecology Research Group, University of Natal, Durban 4041, South Africa Jason D. Hoeksema Department of Environmental Sciences and Policy, 1 Shields Avenue, University of California at Davis, Davis, CA 95616, USA vii viii contributors Jan A. R. A. M. van Hooff Ethologie en Socio-oecologie Groep, Universiteit Utrecht, Pb. 80.086, 3508 TB, Utrecht, The Netherlands Yoh Iwasa Department of Biology, Faculty of Science, Kyushu University, Fukuoka 812-8581, Japan Rebecca J. Lewis Department of Biological Anthropology and Anatomy, Box 90383, Duke University, Durham, NC 27708-0383, USA Ronald Noë Lab. Ethologie des Primates, CEPE–CNRS 2130, 7 rue de l’Université, F-67000 Strasbourg, France and Max-Planck-Institut für Verhaltensphysiologie, Seewiesen, Starnberg, Germany Charles L. Nunn Department of Biology, 221 Gilmer Hall, University of Virginia, Charlottesville, VA 22903-2477, USA Elinor Ostrom Center for the Study of Institutions, Population and Environmental Change, Indiana University, 513 North Park Bloomington, IN 47408- 3895, USA Geoffrey A. Parker Population and Evolutionary Biology Research Group, School of Biological Sciences, University of Liverpool, Liverpool L69 3GS, UK Bogusl- aw Pawl- owski Department of Anthropology, University of Wroclaw, ul. Kuznicza 35, 50-138 Wroclaw, Poland Andrew Pomiankowski Department of Biology, University College London, 4 Stephenson Way, London NW1 2HE, UK Mark W. Schwartz Department of Environmental Science and Policy, 1 Shields Avenue, University of California at Davis, Davis, CA 95616, USA Preface The constraints of life force every organism to behave economically. The study of economic behaviour is therefore not limited to ‘economics’. Scientific disciplines that concentrate on other aspects of human behav- iour, such as psychology, sociology and anthropology, have to pay close attention to the economic decisions that drive human behaviour. The same is true for biological disciplines in which strategic options of indi- vidual organisms play a central role, notably ethology, behavioural ecology and evolutionary ecology. In order to determine in how far a behavioural strategy is ‘economical’, it has to be compared to some ideal norm: the strategy that would yield maximum payoff. In economics this norm is set by the strategy of a hypothetical ‘super-rational’ individual. For most of us the term ‘super-rational’ has a connotation of ‘very intel- ligent’; a strategy achieved by the use of superior cognitive capacities. Evolutionary biologists have realised, however, that even the dimmest of organisms, such as fungi and flatworms, often use ‘super-rational strate- gies’, because a process of selection running over a vast number of gen- erations can shape the behaviour of every species to near perfection, as long as the circumstances in which it lives change slow enough com- pared to the rate of evolution. There lies the common ground between biology and economics: while Adam Smith’s human producers and con- sumers are driven by the ‘invisible hand’ of self-interest, Charles Darwin’s living organisms are driven by the selection for maximising individual fitness. The present volume gives a number of examples of the common grounds on which economics, biology and the social sciences meet. In his introductory chapter Peter Hammerstein shows that there are striking parallels between the use of game theory in economics and evolutionary ix