Social Behaviour in Mammals
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1 TERTIARY LEVEL BIOLOGY Social Behaviour in Mammals Trevor B. Poole, B.Sc., Ph.D. Blackie Glasgow and London Distributed in the USA by Chapman and Hall New York
2 Blackie & Son Limited Bishopbriggs Glasgow G64 2NZ Furnival House High Holborn London WC1 V 6BX Distributed in the USA by Chapman and Hall in association with Methuen, Inc. 733 Third Avenue, New York, N.Y Blackie & Son Ltd First published 1985 All rights reserved No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, recording or otherwise, without prior permission of the Publishers British Library Cataloguing in Publication Data Poole, Trevor Social behaviour in mammals.-(tertiary level biology) 1. Mammals-Behavior 2. Social behavior in animals I. Title II. Series QL739.3 ISBN-13: e-isbn-13: om: / For the USA, International Standard Book Number is X (pbk) Thomson Litho Ltd, East Kilbride, Scotland
3 Preface Mammalian sociobiology is a rapidly advancing field which has made enormous strides in the last ten years. The last major monograph on the subject (Ewer, 1968) was published sixteen years ago, and there is a need for this information to be examined in terms of modern sociobiological theory. My approach throughout is evolutionary and is therefore directed strongly towards research which throws light on the ways in which mammals behave in their natural environments. I have tried to cover as wide a range of mammalian species as possible, although, in some cases, the only data available were obtained from captive individuals. The coverage of this book is not a reflection of the volume of literature published on different species, as I have tried to avoid undue emphasis on the social behaviour of primates and laboratory rodents. I have made scrupulous efforts throughout to avoid an anthropomorphic approach to mammalian behaviour. Terms such as 'strategy', 'evaluation' or 'choice' do not therefore imply conscious planning, but are used neutrally in the way in which they would be applied to a chess-playing computer. In the case of mammals, the programmer was natural selection. While I am fully aware that human beings are mammals, any detailed consideration of human social behaviour lies outside the scope of this book. However, the book may provide a complementary text to those interested in that subject. A glossary of technical terms likely to be unfamiliar to general readers is included towards the back of the book, together with an outline classification of the Mammalia which covers those species mentioned in the text. I am grateful to a number of people for helpful and stimulating discussions. They include Pat Bateson, Brian Bertram, Warren Brockelman. Alan Dixson. Robin Dunbar. Sian Evans, Peter Gortvai. Keith Kendricks, Morris Gosling and Bob Martin, but I take full responsibility for the views I have expressed. I must also thank Pat Bateson, Robin Dunbar, Morris Gosling and Miranda Stevenson for allowing me to quote from review papers in the press. iii
4 iv PREFACE I must express my appreciation to the University College of Wales for granting me study leave, which enabled me to visit other institutions, both to take advantage oflibrary facilities and to discuss the subject matter with other workers in the field. I am particularly grateful to Pat Bateson for providing me with facilities to work at the Sub-Department of Animal Behaviour, University of Cambridge, and to John Hearn and the Zoological Society of London for awarding me an Honorary Research Fellowship which enabled me to complete my writing under ideal conditions. Last, but not least, I must thank Helen for encouraging me to write this book and for bearing with me during its two-year gestation period. T. B. P.
5 Contents Chapter 1. INTRODUCTION 1.1 Mammalian classification I 1.2 Social communication Altruism and inclusive fitness Determinants of mammalian social organization Summary 8 Chapter 2. SENSES AND SOCIAL COMMUNICATION Chemical communication II Perception and production of odours The use of odours in social communication Acoustic communication Sound production The messages conveyed by sounds Visual communication Head Body Tail Tactile communication Summary 38 Chapter 3. REPRODUCTIVE BEHAVIOUR Sexual behaviour 40 3.l.l Theoretical considerations Mating Factors influencing mate choice Inbreeding avoidance and dispersal Adult-young interactions General attributes Lactation Maternal care and developmental needs of the young Paternal care Alloparental behaviour Summary 80 v
6 vi CONTENTS Chapter 4. COMPETITIVE AND AFFILIA TIVE BEHAVIOUR Agonistic behaviour Weapons and fighting techniques The role of aggressive displays Submissive and defensive displays Agonistic displays and their underlying motivation Dominance hierarchy Infanticide Social play A definition of play Forms taken by social play The function of social play Affectional behaviour Summary 118 Chapter 5. SOCIOECOLOGY 120 Chapter Social determinants Types of mammalian social organization The ecology of social organization Diet and food distribution Climate as a determinant of social structure Primate social organization Territorial behaviour Economics of territorial defence Territoriality: mammalian examples Summary 154 AN ORDER-BY-ORDER SYNOPSIS OF SOCIAL BEHAVIOUR Monotremata Marsupialia Insectivora Macroscelidea (elephant shrews) Scandentia (tree shrews) Pholidota (pangolins) Edentata Lagomorpha (rabbits, hares and pikas) Rodentia Hyracoidea Tubulidentata (aardvarks) Perissodactyla (odd-toed ungulates) Proboscoidea (elephants) Artiodactyla (even-toed ungulates) Carnivora Fissipedia (terrestrial carnivores) Pinnipedia (seals. sea lions and walruses) Sirenia (sea cows) 174
7 Chapter 7. CONTENTS 6.17 Cetacea (whales) Mysteceti Odontoceti 6.18 Chiroptera 6.19 Derrnoptera (galagos or flying lemurs) 6.20 Primates Prosimians (suborder Strepsirhini or Prosimii) Simians (suborder Haplorhini) Hominidae 6.21 Summary FUTURE DEVELOPMENTS vii APPENDIX Table I: Species of mammals mentioned in the text classified into orders Table 2: Glossary BIBLIOGRAPHY INDEX
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