Observing Animal Behaviour This Book Is Dedicated to the Memory of Niko Tinbergen (1907–1988) Observing Animal Behaviour

Observing Animal Behaviour This book is dedicated to the memory of Niko Tinbergen (1907–1988) Observing Animal Behaviour Design and analysis of quantitative data Marian Stamp Dawkins Mary Snow Fellow in Biological Sciences Somerville College Oxford and Animal Behaviour Research Group Department of Zoology University of Oxford 1 3 Great Clarendon Street, Oxford OX2 6DP Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc., New York © Marian Stamp Dawkins 2007 The moral rights of the author have been asserted Database right Oxford University Press (maker) First published 2007 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, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this book in any other binding or cover and you must impose the same condition on any acquirer British Library Cataloguing in Publication Data Data available Library of Congress Cataloging in Publication Data Data available Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India Printed in Great Britain on acid-free paper by Biddles Ltd., King’s Lynn ISBN 978–0–19–856935–0 978–0–19–856936–7 (Pbk.) 10987654321 Contents PREFACE vii 1 The power of observation 1 2 Asking the right question 14 3 When all you need is one 43 4 Three principles of observational design 52 5 The selective observer 72 6 Down to detail 89 7 Observing in farms, in zoos, and in the wild 110 8 Analysing observations 116 9 Further observations 125 10 Observing the future 136 REFERENCES 146 Appendix 1. Some random numbers 151 Appendix 2. Power and sample size 152 Appendix 3. Beaufort wind scale 153 INDEX 155 This page intentionally left blank Preface There are several different reasons for writing a book on observing animal behaviour. The first reason (the one that drove me, with a considerable degree of frustration,to think about writing this book in the first place) is the tendency of students and colleagues to ignore the observational phase of a behavioural study and go straight into designing complex experiments without really understanding their animals at all. This often leads them to overlook important aspects of the behaviour of a species, sometimes to design completely inappropriate or impractical experiments, and then to be frustrated when the experiment doesn’t ‘work’.This book aims to show that a little time spent observing goes a long way to improving research, even if it may also, in the end, involve experimentation. The second reason grew out of my own interest in animal welfare, particularly that of farm animals. There is a great deal of interest in the welfare of animals in zoos, on farms and in the wild, all places where experiments may not be possible or ethically desirable. This is where observation really comes into its own, particularly where it is systematic and done on a large scale. Animals can be studied ‘in situ’, in the place a where they live, and so any results are directly applicable to those situations. The replication needed for statistical analysis can often be achieved by combining data from different sources, giving a picture of the ‘real world’ that tightly controlled experiments in one place cannot do.The potential of observation as a non-invasive research tool has not yet been fully appreciated and one of the aims of the book is to show people just what can be accomplished without the need for any sort of experimental manipulation. This leads to a third reason for an interest in observation. The advent of CCTV,video, GPS tracking and other devices that can be installed in farms, zoos,and even on wild animals themselves,now allows us to study the behaviour of undisturbed animals in places where we could not previously follow them and for longer and in more detail than was ever possible before. No longer does observation just mean that which can be collected with pen and paper (although pen and paper still holds an honourable place in behavioural observation). Observation—taken broadly to include the new technology— has an immense potential in the study of behaviour. The aim of this book is viii PREFACE therefore to introduce students (and others) to the power of observation, starting with simple and easily accessible methods suitable for student projects, but opening their eyes to the possibilities that now exist for much more sophisticated analysis of observational data. It aims to show that observational techniques are not the poor relation of experiment but hugely important and informative in their own right. I should perhaps say what this book is not. It is not a statistics textbook, although I would like to think that at least some readers will come to a better understanding of what statistics is all about in a very painless way through reading about animal behaviour.It should be read alongside one of the excel- lent introductory books on statistics that are now available, such Grafen and Hails’ Modern Statistics for the Life Sciences (2002). Nor is it intended as a comprehensive account of methodology in behavioural research. Martin and Bateson’s Measuring Behaviour (1993) and Paul Lehner’s Handbook of Ethological Methods (1996) both go into considerably more detail on techniques, methods, and different experimental designs and should be con- sulted for a more advanced treatment of topics covered here. Bonnie Ploger and Ken Yasukawa’s Exploring Animal Behavior in Laboratory and Field (2003) provides many more specific examples and ideas for actual projects. This book’s role is as a companion, a guide to the study of animal behaviour. It is primarily intended for students doing undertaking their first research project but I hope it will also be useful to anyone who as ever been fascinated by animal behaviour and wanted to know how to study it further. It stresses the supreme importance of asking the right questions but does not pretend to give a detailed list of all the answers. On the contrary, it aims to get people to think for themselves about what the whole process of doing research is all about and how they might go about finding their own answers in one of the most absorbing areas in the whole of biology. It emphasizes observation rather than experimentation, but observation that has gone beyond ‘just’ watching to a science in its own right. I would like to thank Robin McCleery and two anonymous referees for their helpful comments on an early draft of the manuscript. Generous friends and colleagues allowed me to use their wonderful photographs. SBC provided inspiration I could not have done without. 1 The power of observation Imagine it is a bright sunny day and that you are sitting on the wall of a harbour with your back leaning against a warm stone wall, idly watching some gulls poking around in the sandy mud below you. The sun glances off the sea in the distance and even the mud,left wet and glistening by the ebbing tide, has a beauty of its own. The warmth of the wall against your body and the sun on your face are soporific and you are almost half asleep when you are suddenly jolted out of your reverie by the most extraordinary sight. The gull nearest to you is paddling its feet rapidly up and down as it if were running furiously on the spot. As you watch, you realise that this high speed pummelling takes it ever so slightly backwards and that every few seconds it pauses, pecks the mud and apparently finds something to eat.What on earth can it be doing? And it is not just one gull. Several others are taking up the same peculiar behaviour and there are splashy noises made by pairs of vibrating webbed feet all round you.Why? What makes them do it? What do they get out of it? Why haven’t you seen this behaviour before? Is there something special about this day or this state of the tide? Are they following each other’s example? Does it really help them to find food? If so, how? Your idle reverie is now shattered. Your apparently casual observations could be about turn to you into a detective and set you on the trail of asking some very illuminating questions about the behaviour of animals. However, while it is obvious that your observations of what animals do has led you to ask questions about their behaviour, it may be less obvious that further observations, ideally more systematic and quantitative than your original ones, are also crucial to finding the answers. 1.1 Testing hypotheses Many people make the mistake of thinking that science just means perform- ing experiments, as though you couldn’t get anywhere, at least not anywhere scientifically respectable, without actively manipulating the world in some way.

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