Modelling in Behavioural Ecology: An Introductory Text PDF
Preview Modelling in Behavioural Ecology: An Introductory Text
VVeymouth College Modelling in Behavioural Ecology Studies in Behavioural Adaptation Series Editor: John Lazarus, Department of Psychology, The University of Newcastle upon Tyne Gulls and PIOfJers: The Ecology and Behaviour ofM ixed Species Feeding Groups C.]. Barnard and D.B.A. Thompson (Croom Helm, London & Sydney/Columbia University Press, New York) Modelling in Behavioural Ecology: An Introductory Text DENNIS LENDREM, Department of Psychology, University of Newcastle upon Tyne CROOM HELM London & Sydney TIMBER PRESS Portland, Oregon © 1986 Dennis Lendrem Croom Helm Ltd, Provident House, Burrell Row, Beckenham, Kent BR3 lAT Croom Helm, Australia Pty Ltd, Suite 4, 6th Floor, 64-76 Kippax Street, Surry Hills, NSW 2010, Australia British Library Cataloguing in Publication Data Lendrem, Dennis Modelling in behavioural ecology: an introductory text.-(Studies in behavioural adaptation) 1. Ecology-Mathematical models I. Title II. Series 574.5'0724 QH541.l5.M3 ISBN-13: 978-0-7099-4119-4 e-ISBN-13: 978-94-011-6568-6 DOl: 10.1007/978-94-011-6568-6 First published in the USA 1986 ;by Timber Press, 9999 S.W. Wilshire, Portland, OR 97225, USA All rights reserved ISBN-13: 978-0-7099-4119-4 Contents Series Editor's Foreword Vll Acknowledgements x Chapter 1. Introduction 1 Chapter 2. Mathematical Methods 7 Chapter 3. Optimising a Single Behaviour 1: Optimal Foraging Theory 35 Chapter 4. Optimising a Single Behaviour 2: Stochastic Models of Foraging Behaviour 58 Chapter 5. Temporal Patterns: Vigilance in Birds 83 Chapter 6. Behaviour Sequences: Feeding and Vigilance 103 Chapter 7. Short-term and Long-term Optimality Models: Territoriality 123 Chapter 8. Games Theory Models: Social Behaviour 162 Chapter 9. Conclusions 196 Appendix 1: Greek Symbols 205 Appendix 2: Indices 205 References 207 Subject Index 211 Species Index 214 Series Editor's Foreword In the early years of this century a Scottish doctor speculated on the evolutionary origin of human tears. It seemed to him that with the increase in brain size and cognitive powers of our early ancestors many events in the struggle for existence would be just too distressing to observe. How comforting then, for the mother, distraught by the sight of her child being devoured by a lion, to cloud her vision with a flood of tears! Just so, though if the good doctor had pondered further, the following picture might have occurred to him, comfortable in his speculative armchair, and given him some pause for thought. ~e ~, ~. ~ ~'I" .. ~.. - ..... ~ VII viii SERIES EDITOR'S FOREWORD These stories do not, of course, get us very far in explaining the evolution of tears or anything else, but they do remind us how far the study of beha vioural adaptation has come this century. This is, in fact, an exciting time for students of behaviour. The last twenty years have seen a great advance in the theoretical armoury for tackling problems of behavioural evolution and adaptation, and a parallel expansion in empirical studies, particularly in the field. The concepts of inclusive fitness and evolutionary stability, for example, have helped to explain major features of social behaviour and have generated entirely new questions and predictions for the field worker to examine. Cost benefit analysis and optimisation theory have done the same for behaviour in general, and links with population biology and population genetics are becoming stronger. The heady days which saw the birth of behavioural ecology and socio biology are now over, the new concepts have been refined and consolidated, and field data and comparative studies have accumulated at an impressive rate. Now seems a good time to take stock, to review the state of the art and to point some directions for future work. These are the aims of the present series, which will examine questions of behavioural adaptation and evolution in depth. As for our intended readership, we hope that all those interested in such problems, from advanced undergraduate to research worker and lecturer, will find these books of value. Some contributions to the series will encompass particular areas of study, reviewing theory and data and present ing fresh ideas. Others will report the findings of new empirical studies of an extensive nature, which make a significant contribution by examining a range of interrelated questions. The richness, but also the difficulty, of functional enquiry results from the multiple effects of selection pressures and the com plex causal relationships between the behavioural responses to evolutionary forces. Studies which measure a comprehensive set of behavioural attributes, and their ecological correlates, are therefore particularly valuable. The present book holds a special place in the series since it deals with the methods of modelling in behavioural ecology, methods that are common to all areas of the subject. For any behavioural problem it is possible to develop a model that predicts the strategy that maximises the individual's fitness, or some more immediate consequence of its actions. Since natural selection favours behaviour that increases fitness, such models allow us to see how far evolutionary forces have gone towards optimising the individual's answer to a particular behavioural problem, be it feeding, mating, territory size or what ever. In addition, when the success of a behavioural strategy is frequency dependent, evolutionarily stable strategy theory can be employed to predict the outcome. These methods now play a central part in the development of all branches of behavioural ecology and research workers are consequently finding it increasingly important to learn these methods in order to develop their own models and to understand those of their colleagues. Students, too, can benefit SERIES EDITOR'S FOREWORD IX greatly from a working knowledge of these techniques, both for their intrinsic importance to the subject and as a creative aid to understanding the essentials of a problem, and the inevitable assumptions and simplifications required if a solution is to be formulated. In the pages that follow, Dennis Lendrem sets out the general techniques of behavioural modelling, taking great pains to explain the mathematical principles fully so that even the novice can use this book as a primer in the subject. His method is to use published models to illustrate more general behavioural problems. Drawing on discussions with many of the modellers concerned he is also able to illuminate the process by which a model is developed, as well as explaining the nature of the finished product. By work ing with both examples and general principles it is hoped that intending modellers will be able to use this book to transform their ideas into quanti tative and testable models. John Lazarus Newcastle upon Tyne
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