Social Behaviour in Farm Animals
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Color profile: Disabled Composite Default screen SOCIAL BEHAVIOUR IN FARM ANIMALS 1 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:01 A3952:AMA:Keeling:Second Revise:13-Mar-01 Color profile: Disabled Composite Default screen 2 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:01 A3952:AMA:Keeling:Second Revise:13-Mar-01 Color profile: Disabled Composite Default screen Social Behaviour in Farm Animals Edited by L.J. Keeling Department of Animal Environment and Health Swedish University of Agricultural Sciences Skara Sweden and H.W. Gonyou Prairie Swine Centre Saskatoon Canada CABI Publishing 3 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:02 A3952:AMA:Keeling:Second Revise:13-Mar-01 Color profile: Disabled Composite Default screen CABI Publishing is a division of CAB International CABI Publishing CABI Publishing CAB International 10 E 40th Street Wallingford Suite 3203 Oxon OX10 8DE New York, NY 10016 UK USA Tel: +44 (0)1491 832111 Tel: +1 212 481 7018 Fax: +44 (0)1491 833508 Fax: +1 212 686 7993 Email: [email protected] Email: [email protected] Web site: http://www.cabi.org © CAB International 2001. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Social behavior in farm animals / edited by L.K. Keeling and H.W. Gonyou. p. cm. Includes bibliographical references (p. ). ISBN 0-85199-387-4 (alk. paper) 1. Domestic animals--Behaviour. 2. Social behavior in animals. I. Keeling, L.K. (Linda K.) II. Gonyou, H.W. (Harold W.) SF756.7.S58 2001 636′.001′5915--dc21 00-056479 ISBN 0 85199 387 4 Typeset by AMA DataSet Ltd, UK. Printed and bound in the UK by Biddles Ltd, Guildford and King’s Lynn. 4 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:04 A3952:AMA:Keeling:Second Revise:13-Mar-01 Color profile: Disabled Composite Default screen Contents Contents Contributors xiii Acknowledgements xvii Introduction 1 References 4 Part I: Concepts in Social Behaviour 5 1 Living in Groups: an Evolutionary Perspective 7 Michael Mendl and Suzanne Held 1.1 Introduction 8 1.2 Basic Principles: Natural Selection and Group Life 9 1.2.1 Natural selection and behaviour 9 1.2.2 Kin selection and cooperation between relatives 11 1.2.3 Cooperation between non-relatives 12 1.2.4 Studying past and present benefits and costs of group life 13 1.3 Feeding Without Being Fed Upon: Benefits and Costs of Group Life 14 1.3.1 Foraging advantages of group living 15 1.3.2 Avoiding predators: detection, dilution and defence 16 1.3.3 Thermal benefits 17 v 5 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:05 A3952:AMA:Keeling:Second Revise:13-Mar-01 Contents Color profile: Disabled Composite Default screen vi Contents 1.3.4 Disadvantages of group living: competition, conspicuousness, contamination and cuckoldry 17 1.3.5 Confusing the consequences of group living with selection pressures – and other pitfalls 18 1.4 Group Size and the Dynamic Nature of Groups 19 1.4.1 Resource distribution: ideal free distribution theory 19 1.4.2 Optimal group size 20 1.4.3 Resource distribution and defence 21 1.5 Within-group Dynamics: Effects of Dominance 21 1.5.1 Competition or cooperation between dominant and subordinate group members 22 1.6 Group Structure: the Influence of Mating Systems 24 1.6.1 Sex differences and parental care as determinants of mating systems 24 1.6.2 Resource distribution and types of mating system 25 1.6.3 Mating systems and offspring emigration 26 1.6.4 Seasonal variation in group structure 26 1.7 Conclusions: What Use is the Evolutionary Approach for Understanding Farm Animal Social Behaviour? 27 1.7.1 General principles may guide husbandry design and practice 27 1.7.2 Species-specific knowledge and husbandry design 28 1.7.3 Limitations of the evolutionary approach 30 References 31 2 Group Life 37 A. Cecilia Lindberg 2.1 Introduction 37 2.2 Maintaining the Group 39 2.2.1 Dominance 39 2.2.2 Aggression/threatening 42 2.2.3 Stability of hierarchy 44 2.2.4 Social requirements for dominance order to function 45 2.2.5 Affiliative behaviour 47 2.3 Communication Within the Group 47 2.4 Roles Within the Group 49 2.4.1 Strategies and models 49 2.4.2 Proactive and reactive individuals 51 2.5 Group Size 52 References 54 6 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:05 A3952:AMA:Keeling:Second Revise:13-Mar-01 Contents Color profile: Disabled Composite Default screen Contents vii 3 Parental Behaviour 59 Per Jensen 3.1 Introduction 60 3.2 Parent–Offspring Conflict 60 3.3 Varieties of Parental Behaviour 61 3.4 Early Parental Care 63 3.5 Mother–Offspring Bonds 64 3.6 Types of Food Provisioning 66 3.7 Communication During Suckling 67 3.8 Regurgitation 69 3.9 Begging 69 3.10 Factors Affecting Parental Care 72 3.11 The Transition to Offspring Independence 73 References 76 4 The Evolution and Domestication of Social Behaviour 83 W. Ray Stricklin 4.1 Introduction and Overview 84 4.2 A Brief History of Time – Relative to Domestication and Modern Farm Animals 85 4.3 A Cursory Overview of the Evolutionary Basis of Social Behaviour 89 4.4 The Role of Behaviour in Domestication 92 4.5 Domestication as an Adaptive Trait Leading to Increased Genetic Fitness 95 4.6 Is Domestication a Single-gene Trait? 98 4.6.1 The argument for gradualism 99 4.6.2 The argument against gradualism 100 4.6.3 Possible molecular basis for a gene related to domestication 103 4.7 Impact of Production Trait Selection on Social Behaviour 104 4.8 Beyond Domestication? 104 Notes 105 References 107 Part II: The Social Behaviour of Domestic Species 111 5 The Social Behaviour of Cattle 113 Marie-France Bouissou, Alain Boissy, Pierre Le Neindre and Isabelle Veissier 5.1 Basic Social Characteristics 113 5.1.1 Composition and structure of social groups 114 5.1.2 Use of space 115 7 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:05 A3952:AMA:Keeling:Second Revise:13-Mar-01 Contents Color profile: Disabled Composite Default screen viii Contents 5.1.3 Communicaton 115 5.1.4 Intra-group interactions 117 5.2 Social Behaviour Under Commercial Conditions 120 5.2.1 Social groupings 120 5.2.2 Social structure 121 5.2.3 Effects of group size and space allowance on social behaviour 129 5.3 Social Behaviour, Management and Welfare 130 5.3.1 Influence of social partners 130 5.3.2 Grouping animals 131 5.3.3 Separation problems 131 5.3.4 Dominance-related problems 132 5.3.5 Abnormal behaviour 133 References 135 6 The Social Behaviour of Pigs 147 Harold W. Gonyou 6.1 Basic Social Characteristics 147 6.1.1 Composition and structure of social groups 148 6.1.2 Use of space 149 6.1.3 Communication 149 6.1.4 Cohesion and dispersion 150 6.1.5 Inter-group interactions 151 6.1.6 Intra-group interactions 151 6.2 Social Groupings Under Commercial Conditions 154 6.2.1 Social groupings 154 6.2.2 Social effects of production 160 6.2.3 Effects of group size and space allowance on social behaviour 163 6.3 Social Behaviour, Management and Welfare 165 6.3.1 Grouping animals 165 6.3.2 Separation problems 167 6.3.3 Abnormal behaviour 168 References 168 7 The Social Behaviour of Domestic Birds 177 Joy Mench and Linda Keeling 7.1 Basic Social Structure 177 7.1.1 Composition and structure of social groups 178 7.1.2 Use of space 180 7.1.3 Communication 181 7.1.4 Cohesion and dispersion 182 7.1.5 Inter-group interactions 183 7.1.6 Intra-group interactions 183 8 Z:\Customer\CABI\A3952 - Keeling - Social Behaviour DA #8.vp 13 March 2001 11:04:06 A3952:AMA:Keeling:Second Revise:13-Mar-01 Contents Color profile: Disabled Composite Default screen Contents ix 7.2 Social Behaviour Under Commercial Conditions 187 7.2.1 Social groupings 187 7.2.2 Social effects on production 191 7.2.3 Effects of group size and space allowances on social behaviour 193 7.3 Social Effects on Behaviour, Welfare and Production 195 7.3.1 Grouping animals 195 7.3.2 Separation problems 196 7.3.3 Dominance-related problems 197 7.3.4 Abnormal behaviour 197 References 199 8 The Social Behaviour of Sheep 211 Andrew Fisher and Lindsay Matthews 8.1 Basic Social Structure 211 8.1.1 Composition and structure of social groups 212 8.1.2 Use of space 215 8.1.3 Communication 216 8.1.4 Cohesion and dispersion 218 8.1.5 Inter-group interactions 220 8.1.6 Intra-group interactions 221 8.2 Social Behaviour Under Commercial Conditions 230 8.2.1 Social groupings 230 8.2.2 Effects of group size and space allowance on social behaviour 232 8.2.3 Social effects on production 234 8.3 Social Behaviour, Management and Welfare 236 8.3.1 Grouping and separation problems 236 8.3.2 Social isolation and facilitation 237 8.3.3 Dominance-related problems 238 8.3.4 Abnormal behaviour 238 References 239 9 The Social Behaviour of Horses 247 Natalie K.