CURRENT PROBLEMS IN SOCIOBIOLOGY

CURRENT PROBLEMS IN SOCIOBIOLOGY

CURRENT PROBLEMS IN SOCIOBIOLOGY EDITED BY KING'S COLLEGE SOCIOBIOLOGY GROUP, CAMBRIDGE CAMBRIDGE UNIVERSITY PRESS Cambridge London New York New Rochelle Sydney Melbourne

cambridge university press Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo, Delhi, Dubai, Tokyo, Mexico City Cambridge University Press The Edinburgh Building, Cambridge cb2 8ru, UK Published in the United States of America by Cambridge University Press, New York Information on this title: /9780521285209 Cambridge University Press 1982 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 1982 Re-issued 2010 A catalogue record for this publication is available from the British Library British Library Cataloguing in Puhlication Data Current problems in sociobiology. I. Social behavior in animals Addresses, essays, lectures I. King s College Sociobiology Group 591.5 QL775 isbn 978-0-521-24203-5 Hardback isbn 978-0-521-28520-9 Paperback Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

Contents List of participants Preface by P.P.G. BATESON Acknowledgments Introduction by J. MAYNARD SMITH Vll IX Xlll I Natural selection and sociobiology (edited by R. w. WRANGHAM) 5 R. I. M. DUNBAR Adaptation, fitness and the evolutionary tautology 9 2 J. MAYNARD SMITH The evolution of social behaviour - a classification of models 29 3 R. DAWKINS Replicators and vehicles 45 4 P. O'DONALD The concept of fitness in population genetics and sociobiology 65 II Complexity in evolutionary processes ( edited by D. I. RUBENSTEIN) 87 5 D.1. RUBENSTEIN Risk, uncertainty and evolutionary strategies 91 6 E.A. THOMPSON Gene competition without selection 113 7 P.P.G. BATESON Behavioural development and evolutionary processes 133 8 A. LOMNICKI Individual heterogeneity and population regulation 153

III Evolutionaryconflictsofinterest(editedbYR.I.M. DUNBAR) 169 9 G.A. PARKER Phenotype-limited evolutionarily stable strategies 173 10 N. KNOWLTON Parental care and sex role reversal 203 11 T.H. CLUTTON-BROCK AND S.D. ALBON Parental investment in male and female offspring in mammals 223 IV Sociality (edited by B.C.R. BERTRAM) 249 12 B.C.R. BERTRAM Problems with altruism 251 13 R.W. WRANGHAM Mutualism, kinship and social evolution 269 14 N. A. CH AG NON Sociodemographic attributes of nepotism in tribal populations: man the rule-breaker 291 V The problems of comparison (edited by T.H. CLUTTON- BROCK) 319 15 P.J. JARMAN Prospects for interspecific comparison in sociobiology 323 16 P.H. HARVEY AND G.M. MACE Comparisons between taxa and adaptive trends: problems of methodology 343 17 N.B. DAVIES Behaviour and competition for scarce resources 363 Index 381 vi

List of Participants S.D. ALBON, Research Centre, King's College, Cambridge, England, and Department of Zoology, Large Animal Research Group, 34A Storeys Way, Cambridge, England P.P.G. BATESON, Sub-Department of Animal Behaviour, University of Cambridge, Madingley, Cambridge, England B.C.R. BERTRAM, Research Centre, King's College, Cambridge, England and Zoological Society of London, Regent's Park, London, NW1 4RY, England N.A. CHAGNON, Department of Anthropology, Pennsylvania State University, University Park, Pennsylvania 16802, USA T.H. CLUTTON-BROCK, Research Centre, King's College, Cambridge, England and Department of Zoology, Large Animal Research Group, 34A Storeys Way, Cambridge, England N. B. DAVIES, Department of Zoology, University of Cambridge, Downing Street, Cambridge, England R. DAWKINS, Department of Zoology, University of Oxford, South Parks Road, Oxford, England R. I. M. DUNBAR, Research Centre, King's College, Cambridge, England and Sub-Department of Animal Behaviour, University of Cambridge, Madingley, Cambridge, England P. H. HARVEY, School of Biological Sciences, University of Sussex, Falmer, Brighton, Sussex, England P.J. JARMAN, School of Natural Resources, University of New England, Armidale, New South Wales 2351, Australia

N. KNOWLTON, Department of Biology, Yale University, P.O. Box 6666, New Haven, Connecticutt 06520, USA A. LOMNICKI, Institute of Environmental Biology, Jagiellonian University, ul. Karasia 6, 30-060 Krakow, Poland G.A. MACE, Department of Zoology, University of Newcastle upon Tyne, England 1. MAYNARD SMITH, School of Biological Sciences, University of Sussex, Falmer, Brighton, Sussex, England P. O'DONALD, Department of Genetics, University of Cambridge, Downing Street, Cambridge, England G.A. PARKER, Department of Zoology, University of Liverpool, Brownlow Street, Liverpool, Lancashire, England D.1. RUBENSTEIN, Research Centre, King's College, Cambridge, England and Department of Biology, Princeton University, Princeton, New Jersey 08540, USA E.A. THOMPSON, Department of Mathematical Statistics, University of Cambridge, Mill Lane, Cambridge, England R.W. WRANGHAM, Research Centre, King's College, Cambridge, England and c/o Dr I. Devore, Department of Anthropology, Harvard University, Boston, Mass. USA viii

Preface The papers in this volume were originally presented at a conference held at King's College, Cambridge on 4-6 July 1980, to mark the culmination of the King's College Sociobiology project. For this conference, all those who had been closely involved with the project were invited to submit papers concerned with unsolved problems in sociobiology. In order to understand how the King's Sociobiology group came to be formed it is necessary to understand that Cambridge Colleges are relatively independent of the University and some have great wealth of their own. Every so often a college that is rich enough can behave like a Renaissance patron and do something that a British university, largely dependent on State support, could rarely do. My own College, King's, decided in the midsixties to use some of its considerable resources to help subjects that looked exciting but did not have an adequate toehold within the University. It established a Research Centre in which a group of people within the same subject could hold Fellowships. Each programme was to last up to five years and, in addition to themselves, members could draw in short-term visitors, research assistants and graduate students. The idea was that a lively group would give sufficient impetus to their subject to keep it going in Cambridge after the project had ended in the Research Centre. Early in 1975 Nick Humphrey and I, as Fellows of King's, suggested to the College that the Research Centre should establish a project in behavioural ecology. During the sixties and early seventies the fruits of numerous field studies of animals were starting to suggest coherent explanations for the ways in which social behaviour might be related to ecological conditions. The ideas were brought together by the increasingly powerful use of evolutionary theory. A mass of seemingly unrelated evidence started to make sense and the subject looked exceptionally promis

ing. King's decided to support behavioural ecology and, once it had done so, the Convenor of the Research Centre, Donald Parry, took over the job of organising the project. Along with running a busy University Department of his own, he handled with consummate efficiency every stage from advertising the Fellowships to administering the most active phase of the programme. A lot of its subsequent success was due to his efforts. After the project had been running for a year, its title was changed from Behavioural Ecology to Sociobiology since we wanted to include functional studies of social behaviour which were not necessarily ecological in character. The term 'Sociobiology' had been in use since the late 1940s; indeed, the present-day American Animal Behavior Society had grown out of a section of the Ecological Society of America and the American Society of Zoologists called 'Animal Behavior and Sociobiology'. However, we should, perhaps, have been quicker to realise how much opinion would be polarized by the recent attempts to inject a particular brand of biology into the social sciences. The lacerations resulting from the ensuing ideological conflict have not yet healed, and in many places 'Sociobiology' is either a battlecry or a term of abuse. Fortunately, the Research Centre project was unaffected, largely because the people who were appointed to the group were strongly committed to empirical research. Nobody who knew their work could accuse them of doing bad science. Furthermore, they would tolerate neither sloppy argument nor extravagant generalisations from studies of animals to humans. After two years of recruiting the Sociobiology Group had four Fellows with longer-term appointments: Brian Bertram, Tim Clutton-Brock, Dan Rubenstein and Richard Wrangham. They were joined by Robin Dunbar who had been awarded one of the much prized Science Research Council Advanced Fellowships. During the course of the project Adam Lomnicki and Geoff Parker each came for one year while on sabbatical leave from their Universities. Peter Jarman, Nancy Knowlton and Napoleon Chagnon worked for shorter periods in the Research Centre and, while they did not hold Fellowships, played vigorous parts in its intellectual life. In addition, Elizabeth Thompson, who was already a Fellow of the College, brought much needed mathematical skills to bear on some of the problems that were being discussed, and Steve Albon collaborated throughout with Tim Clutton-Brock on their study of red deer. Of the numerous other people who, by their talks and discussions, contributed so much to making the years of the project exciting and enjoyable, Nick Davies, Richard Dawkins, Paul Harvey, John Maynard Smith and PeterO'Donald were especially influential. x

One of the most important aspects of the Research Centre's work was the weekly seminar. The talks and the discussions they generated were excellent and, what proved crucial, attracted a regular audience from groups scattered throughout Cambridge and East Anglia. People from different Departments and research groups suddenly found that they had interesting things to say to each other. The Research Centre was operating exactly as it should do and providing a focus for discussion and an impetus for further research. After four productive years which culminated in the conference from which this book has been produced, the Sociobiology Group has now dispersed. What remains behind is a widespread enthusiasm for studies of the evolution of behaviour and behavioural ecology and, at the same time, cool sanity about the issues. That legacy, along with each member's personal contribution to research, is the lasting achievement of the King's Sociobiology Group. October 1980 Patrick Bateson xi

Acknowledgments It is a very real pleasure to thank the Provost and Fellows of King's College for their interest, help and support in the Sociobiology Project over the last five years. For all of us, our time in the Research Centre provided opportunities for research and collaboration which would have been unavailable elsewhere. We are particularly grateful to Pat Bateson and Nick Humphrey for instigating the project; to Donald Parry and Herbert Huppert who administered the Research Centre together with the board of managers, for their help, sympathy and toleration in dealing with our various needs; to Hazel Clarke, the Research Centre's administrative secretary, for secretarial help and unflagging good humour; and to the staff of King's College for their part in making King's a pleasant place to work. We are also grateful to the staff of the Department of Zoology, the Sub-department of Animal Behaviour, and the Computer Centre, all of which provided us with specialized facilities unavailable in the Research Centre. Finally, we should like to thank the large number of colleagues who participated in the Research Centre's activities and whose stimulating interest and encouragement were the spur for so many of our activities. The Editors