The Study of Animal Behaviour

Transcription

1 The Study of Animal Behaviour

2 The Study of Animal Behaviour FELICITY HUNTINGFORD Department oj Zoology University ojglasgotlj, UK LONDON NEW YORK Chapman and Hall

3 Firstpublished 1984 by Chapman and Hall Ltd 11 New Fetter Lane, London EC4P 4EE Published in the USA by Chapman and Hall 733 ThirdAvenue, New YorkNY Felicity Huntingford Photoset by Enset Ltd. Midsomer Norton, Bath, Avon and printed in Great Britain by J. w. Arrowsmith Ltd., Bristol ISBN-13: e-isbn-13: DOl: / This title is available in both hardbound and paperback editions. The paperback edition is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, re-sold, hired out, or otherwise circulated without thepublisher's prior consent in any form of binding or cover other than that in which it is published and without a similar condition including this condition being imposed on the subsequent purchaser. All rights reserved. No part of this book may be reprinted, or reproduced or utilized in any form or by any electronic, mechanical or other means, now known or hereafter invented, including photocopying and rewrding, or in any information storage and retrieval system, without permission in writingfrom the publisher. British Library Cataloguing in Publication Data Huntingford, Felicity A. The study of animal behaviour. 1. Animal behavior I. Title QL751 Library of Congress Cataloging in Publication Data Huntingford, Felicity A. (Felicity Ann) The study of animal behaviour. Bibliography: p. Includes index. 1. Animal behavior. I. Title. QL751.H

4 To Tim and Anne, without whom this book would not have been written, and to Joan and Jessica, without whom it might have been written sooner

5 Contents Preface Acknowledgements List of plates Chapter 1 Introduction 1.1 The ethological approach to the study of behaviour 1.2 A brief outline of classical ethological theory 1.3 The modem study of animal behaviour 1.4 An outline of the book Chapter 2 The description and measurement of behaviour 2.1 Describing behaviour by its function 2.2 Describing behaviour by its form Behaviour described as a continuous process Behaviour described as a series of discrete events 2.3 Describing and measuring the relationship between an animal and its environment Describing the way a single animal uses space Describing the relationship between an animal and neighbouring conspecifics 2.4 "What is the point of all this sophisticated analysis? Chapter 3 The study of the causes of behavioural change 3.1 "What constitutes a causal explanation of behaviour? 3.2 The different kinds of causal explanation 3.3 Motivational models Descriptive and explanatory models General, theory-based and specific, data-based models Deterministic and probabilistic models Assessing models xiv xv xvi Contents vii

6 3.4 Studying external influences on behaviour Prey catching in toads; behavioural studies Attack in Haplochromis burtoni Incubation in herring gulls Prey catching in toads; physiological studies The role of external stimuli in the control of behaviour Studying internal influences on behaviour Identifyin.g groups of causally related acts The role of internal factors in the control of behaviour What is the nature of the mechanisms which cause behavioural change? A probabilistic model A deterministic model Modelling motivation as a hierarchy of decisions Control theory models of motivation Space-state models Studying the physiological bases of behavioural change Physiological explanations of complex behaviour Alteration in behavioural theory as a result of physiological research Identifying fruitful topics for physiological study by behavioural analysis Ethological concepts and techniques can improve the quality of physiological research Ethological ideas can help in the interpretation of physiological results 103 Chapter 4 The development of behaviour Problems with the instinct-learning dichotomy Instincts are defined by exclusion Interpreting deprivation experiments is not simple Learning is not the only way the environment influences development of behaviour The instinct -learning dichotomy confounds three distinct consequences of behavioural development Why is the term innate still used? Describing the ontogeny of behaviour The emergence of recognizable behaviour patterns in young animals The time of appearance of behaviour patterns during development 111 Vlll Contents

7 4.3.3 The development of motivational systems Physiological changes accompanying behavioural development Characterizing the factors which influence the development of behaviour Deductions from descriptive studies Experimental studies of behavioural development Experimental studies of the development of song in passerines A quantitative model of the development of social preferences in birds Classifying the factors which influence the development of behaviour Some general features of behavioural development Sensitive periods Constraints on learning Functional aspects of behavioural development 136 Chapter 5 The adaptive significance of behaviour Sources of evidence about the adaptive significance of behaviour Deduction from the theory of natural selection The context in which behaviour is shown The short-term consequences of behaviour Comparisons between species and populations Measuring the fitness of individuals which differ in their behaviour Experimental manipulation of the consequences of behaviour Deductions from the genetic architecture of behaviour Models of the adaptive significance of behaviour Difficulties in studying the adaptive significance of behaviour Multiple benefits Costs, benefits and optimality models Frequency dependent costs and benefits; games theory The state of the art The adaptive significance of the way animals pattern their behaviour in time Rhythms ofbella!iollr _ The adaptive significance of behavioural rhythmicity The adaptive significance of the way animals use space Dispersal patterns Deciding where to live What to do where; optimal foraging theory Problems with optimal foraging theory 180 Contents ix

8 5.5.5 Constraints on use of space resulting from the presence of conspecifics Territorial behaviour The adaptive significance of an animal's aggressive responses Games theory and the hawk-dove model Weaknesses of the hawk-dove model Games theory models in sexually reproducing species Conflicts between doves; the War of Attrition model Games with progressive escalation Games with asymmetries between opponents Games in which information is acquired during a fight Gamel> with non-random encounters The contribution of games theory to our understanding of the adaptive significance of the way animals fight The adaptive significance of an animal's breeding habits; mating systems Life history parameters Questions about sex Mating systems Adaptive significance of behaviour accompanying mating Identifying the correct species Preventing mating with conspecifics of the wrong sex Synchronizing the behaviour of potential mates The impact of intra-sexual competition for mates The impact of selectiveness in potential mates Alternative mating strategies Darwin, natural selection and sexual selection Conflicts of interest between the sexes The adaptive significance of parental care The adaptive significance of living in groups Studies of the adaptive significance of group life The benefits resulting from group life OJ The complex costs and benefits of group life; Caraco's model (1979) Behaviour which cannot be explained by classic natural selection theory An outline of the problem Animals may ultimately benefit from their 'altruistic' behaviour J Group selection 257 x Contents

9 Kin selection Parental manipulation Has altruism been explained? Overview; sociobiology and behavioural ecology Weaknesses in the sociobiological approach to behaviour Reasons why behavioural differences may not be adaptive 272 Chapter 6 The phylogeny of behaviour Sources of evidence about the phylogeny of behaviour The fossil record Ontogeny The behaviour of interspecific hybrids Comparative studies Some representative behavioural phylogenies Tongue movements in reptiles Predatory behaviour in gastropods Aggressive interactions in fish Hymenopteran social systems Deriving general principles of behavioural evolution Sound production in crickets Courtship in peacocks 288 Chapter 7 The role of behaviour in the evolutionary process The behaviour of other animals as a major selective force The behaviour of conspecifics The behaviour ofheterospecifics Behaviour dictates the selection pressures to which an animal is exposed The impact of behaviour on population structure Behavioural influences on gene flow between populations Behavioural influences on population size Behavioural influences on panmixis 299 Chapter 8 Behavioural genetics The objectives of research into the inheritance of behaviour Potential contributions of genetics to the study of animal behaviour As an additional research tool in the analysis of behaviour As a source of essential information about behaviour Quantitative genetics Techniques for identifying the genetic origin of a known behavioural difference 306 Contents xi

10 8.4.1 Crosses between behavioural variants Selective breeding from extremes of a behavioural continuum Screening known genetic variants for behavioural differences Known single gene mutations which influence behaviour Visible differences in chromosome structure with behavioural effects Comparing naturally occurring, isolated populations Correlating degree of relatedness and behavioural similarity Inbred strains Itecombinantstrains Characterizing the precise behavioural effects of genetic differences The mechanisms whereby genes influence behaviour Effects on the structures necessary for performing the behaviour Effects on muscles or neuromuscular junctions Effects on the sense organs Effects on the CNS Effects on the endocrine system Effects on some totally unknown mechanism Genetic mosaics Animal behaviour and behavioural genetics 333 Chapter 9 Applied ethology Clarification of terms; what is applied ethology? Ways in which ethological research can be applied to practical problems Direct application of the results of ethological research Using the ideas and concepts of ethology Using ethological techniques Pest control Increasing the productivity of commercially important species Identifying suitable species for exploitation Selection for desired behavioural attributes Controlling dispersal and movement Promoting survival Promoting growth Promoting successful reproduction Animal welfare Identifying suffering in animals Alleviating animal suffering Conservation 354 xii Contents

11 9.7 Human behaviour Applying ethological techniques to human behaviour Ethological concepts as a source of hypotheses about human behaviour Making direct use of the facts collected by ethologists Sociobiology and human behaviour 366 References Author index Species index Subject index Contents Xlll

12 Preface The aim of this book is to identify the main areas of active discussion about, and research into, the biology of animal behaviour, to describe and assess ways in which these can be studied and using selected examples, to illustrate the kinds of results which are emerging. It is not intended to provide an exhaustive review of all we know about animal behaviour, although the examples have been chosen to cover as many as possible of the things that animals do. XlV Preface

13 Acknowledgements I would like to thank C. Swann, M.L.N. Murthy and the Superbrain for typing the manuscript; Linda Partridge, Pat Monaghan, Douglas Fraser and Richard Wilson for constructive criticism of earlier drafts; Alan Crowden for help in planning and producing the book and, particularly, Tim Huntingford for help and encouragement at all stages of its production. Acknowledgements are gratefully made to Jim Tulley fot producing plates 1, 2, 3, 7 and 12 and to Michael Hansell for the remainder. Acknowledgements xv

14 List of plates Plate 1 Non-random spacing in animals; a colony of breeding gannets 30 Plate 2 Behaviour of gulls at the nest; relief of an incubating kittiwake by its partner 58 Plate 3 Development of behavioural sequences; certain elements of play in cats become incorporated into predatory sequences 113 Plate 4 Foraging bees 175 Plate 5 Sexual advertisement? A male frog calling at a breeding pond 222 Plate 6 Parental investment by the female; a mother spider monkey carrying her infant 238 Plate 7 Group living and vigilance; a flock of grazing pink-footed geese 246 Plate 8 Living in groups can improve feeding efficiency; social facilitation in pink-footed geese 248 Plate 9 Sterile castes (of termites), a problem for the conventional theory of natural selection 266 Plate 10 Factors other than kinship may favour communal nesting in hymenoptera; a hornet (Vespa tropica) destroying a larva of the wasp Parischnogaster mellyi 269 Plate 11 Possible stages in the evolution of complex social organization in the hymenoptera. (a) A single female Parischnogaster jacobsoni at her nest; (b) a small group of Liostenogastervaripiaa at a communal nest 286 Plate 12 Differential choice of habitat by closely related species; the voles Clethrionomys brittanicus (a) and Microtus agrestis (b) in their natural environments 297 xvi List of plates