... "1, '",.. r.",. r ",~ Illustrated by Leslie M. Downie Department of Zoology, University of Glasgow

Transcription

1 ANIMAL CONFLICT

2 Illustrated by Leslie M. Downie Department of Zoology, University of Glasgow "1,... '",.. r.",. r ",~

3 ANIMAL CONFLICT Felicity A. Huntingford and Angela K. Turner Department of Zoology, University of Glasgow.-~ '... '.. I ->,, f. ~ :"fi London New York CHAPMAN AND HALL

4 Chapman and Hall Animal Behaviour Series SERIES EDITORS D.M. Broom Colleen Macleod Professor of Animal Welfare, University of Cambridge, UK P.W. Colgan Professor of Biology, Queen's University, Canada Detailed studies of behaviour are important in many areas of physiology, psychology, zoology and agriculture. Each volume in this series will provide a concise and readable account of a topic of fundamental importance and current interest in animal behaviour, at a level appropriate for senior undergraduates and research workers. Many facets of the study of animal behaviour will be explored and the topics included will reflect the broad scope of the subject. The major areas to be covered will range from behavioural ecology and sociobiology to general behavioural mechanisms and physiological psychology. Each volume will provide a rigorous and balanced view of the subject although authors will be given the freedom to develop material in their own way.

5 To Tim) Joan and Jessica) with thanks) and to all the children who provided inspiration

6 First published in 1987 by Chapmatl and Hall Ltd 11 New Fetter Lane, London EC4P 4EE Published in the USA by Chapman atld Hall 29 West 35th Street, New York, NY Felicity A. Huntingford and Angela K. Turner Softcover reprint of the hardcover 1 st edition 1987 ISBN -13: This title is available in both hardbound and paperback editions. The paperback edition is sold sll~iect to the condition that it shall 110t, by way of trade or otherwise, be lmt, resold, hired out, or otherwise circulated without the publisher's prior consmt ill ally form o/bindillg or cover other thatl that ill which it is published alld without a similar condition illcluding this collditioll being imposed 011 the subsequmt purchaser. All rights reserved. No part of this book may be reprillted, or reproduced or utilized ill atly form or by any. electronic, mechanical or other means, now known or hereafter invmted, including photocopying and recording, or in any information storage and retrieval system, without permission in writingfrom the publisher. British Library Cataloguing in Publication Data Huntingford, Felicity Animal conflict. 1. Aggressive behaviour in animals I. Title II. Turner, Angela K QL758.5 ISBN-13: Library of Congress Cataloging in Publication Data Huntingford, Felicity. Animal conflict. Bibliography: p. Includes index. 1. Agonistic behaviour in animals. 2. Conflict (Psychology) I. Turner, Angela K., II. Title. III. Title: Animal conflict. QL758.5.H ISBN -13: e-isbn-13: DOl: /

7 Contents Preface Acknowledgements PART ONE PATTERNS OF ANIMAL CONFLICT 1 Conflict in the animal world 1.1 Animal fights 1.2 Conflicts of interest 1.3 Adaptive responses to conflicts of interest 1.4 The problem of definition 1.5 The biological study of animal conflict Xl X11l A survey of animal conflict Introduction Acellular organisms and protists Sea anemones (coelenterates) Parasitic worms (acanthocephalans) Ragworms (annelids) Limpets, octopuses and squids (molluscs) Insects Shrimps and crabs (crustaceans) Scorpions, mites and spiders (arachnids) Sea urchins ( echinoderms) Fish Salamanders and frogs (amphibians) Lizards and snakes (reptiles) Birds (aves) Mammals Issues and concepts in the study of animal conflict How do animals fight? Context - specific agonistic behaviour Alternative ways of winning

8 Vl11 Contents PART TWO THE CAUSES OF AGONISTIC BEHAVIOUR 55 4 Behavioural mechanisms Introduction The systems that control agonistic behaviour The effects of cues from an opponent The effects of other stimuli from the environment The dynamics of undisturbed fights Models of the control of agonistic behaviour Overview 92 5 The role of hormones Introduction Mechanisms Hormones and the patterns of animal conflict Reproductive hormones in vertebrates Adrenal hormones Endogenous opiates Other hormones Invertebrates Overview Neural mechanisms Introduction Neural mechanisms and the patterns of animal conflict Analysing agonistic information Producing agonistic movements Producing changes in responsiveness Effects of fighting experience on brain biochemistry Overview 159 Concluding comments on the causes of agonistic behaviour 162 PART THREE GENETIC AND ENVIRONMENTAL INFLUENCES Genetics 7.1 Introduction 7.2 Genetics and the patterns of animal conflict 7.3 Identifying genetic influences 7.4 Patterns of inheritance 7.5 The route from genes to behaviour 7.6 Overview

9 Contents Development Introduction The agonistic behaviour of very young animals The developmental origin of agonistic movements Playful fighting Changes with age Environmental influences: identifying the relevant factors Environmental influences: kinds of identified effect Environmental influences: the non-social environment Environmental influences: the social environment Overview Genes, environments and agonistic behaviour PART FOUR. CONSEQUENCES, FITNESS AND EVOLUTIONARY CHANGE The consequences of animal conflict Introduction Consequences for individuals Consequences for populations Consequences for ecological communities Evolutionary consequences Overview IX Evolutionary history Introduction Evolutionary origins Evolutionary diversification Overview The behavioural ecology of animal conflict 11.1 Introduction 11.2 Strategies for resolving conflicts 11.3 Costs and benefits of being dominant 11.4 Costs and benefits of territorial behaviour 11.5 Conflict between the sexes 11.6 Conflict within the family 11.7 Infanticide and cannibalism 11.8 Alternative strategies 11.9 Conflict in social groups Overview

10 x The biology of human aggression Introduction What is human aggression? Issues and concepts The causes of aggression in humans Genes, environment and the development of agonistic behaviour The behavioural ecology of human conflict Preventing, predicting and controlling human aggression Overview Contents References Species index Subject index

11 Preface In the past twenty years there have been many new developments in the study of animal behaviour: for example, more sophisticated methods of neurophysiology; more precise techniques for assessing hormonal levels; more accurate methods for studying animals in the wild; and, on the functional side, the growth of behavioural ecology with its use of optimality theory and game theory. In addition, there has been a burgeoning number of studies on a wide range of species. The study of aggression has benefited greatly from these developments; this is reflected in the appearance of a number of specialized texts, both on behavioural ecology and on physiology and genetics. However, these books have often been collections of papers by specialists for specialists. No one book brings together for the nonspecialist all the diverse aspects of aggression, including behavioural ecology, genetics, development, evolution and neurophysiology. Neither has there been a comparative survey dealing with all these aspects. Therefore one of our aims in writing this book was to fill in these gaps. Another of our aims was to put aggression into context with respect to other aspects of an animal's lifestyle and in particular to other ways in which animals deal with conflicts of interest. Aggressive behaviour does not occur in a biological vacuum. It both influences and is influenced by the animal's ecological and social environment, so we consider both the complex antecedent conditions in which aggressive behaviour occurs, and its ramifying consequences in the ecosystem. Throughout the text common names are used wherever they are available. Scientific names are given in the species index.

12 Acknowledgements We are grateful to: Jon Barnes, Pat Bateson, Don Broom, Barry Everitt, John Goss-Custard, Mike Hansell, Alasdair Houston and Marilyn Ramenofsky for constructive and very helpful criticism of early drafts; M. E. Archer, T. Bakker, P. Bateson, P. Bronstein, K. D. Browne, T. Caro, D. A. Collins, D. A. Dewsbury, E. G. Dunning, V. Geist, T. Groothuis, T. R. Halliday, I. R. Inglis, J. Lazarus, P. J. Murphy, E. Nevo, P. W. Sherman, P. Thompson, G. A. Van Oortmersson and J. M. Williams for permission to use unpublished data, for papers seen in press and for recent reprints; P. W. Sherman for assistance with references; Neil Metcalfe for permission to use a photograph and H. Dingle, M. Enquist, Charlie Evans, H. W. Fricke, V. Geist, E. B. Keverne and A. Samuels for use of illustrations; Liz Denton and Peter Rickus for preparation of figures, Clarice Swan and Lynn Mackenzie for help in typing figure legends and tables; and Alan Crowden for being a patient and helpful editor.