THE BEHAVIOUR OF THE DOMESTIC CAT, 2ND EDITION This page intentionally left blank The Behaviour of the Domestic Cat, 2nd Edition John W.S. Bradshaw School of Veterinary Sciences University of Bristol Rachel A. Casey School of Veterinary Sciences University of Bristol Sarah L. Brown School of Veterinary Sciences University of Bristol CABI is a trading name of CAB International CABI CABI Nosworthy Way 38 Chauncey Street Wallingford Suite 1002 Oxfordshire OX10 8DE Boston, MA 02111 UK USA Tel: +44 (0)1491 832111 Tel: +1 800 552 3083 (toll free) Fax: +44 (0)1491 833508 Tel: +1 (0)617 395 4051 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org © J.W.S Bradshaw, R.A. Casey and S.L. Brown 2012. 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 Bradshaw, John W. S. The behaviour of the domestic cat / John W.S. Bradshaw, School of Veterinary Sciences, University of Bristol, Rachel A. Casey, School of Veterinary Sciences, University of Bristol, Sarah L. Brown, School of Veterinary Sciences, University of Bristol. -- Second edition. pages cm Includes bibliographical references and index. ISBN 978-1-84593-992-2 (pbk.) 1. Cats--Behavior. I. Casey, Rachel A. II. Brown, Sarah L. III. Title. SF446.5.B73 2013 636.8--dc23 2012026680 ISBN: 978 1 84593 992 2 Commissioning editor: Rachel Cutts Editorial assistant: Chris Shire Production editor: Tracy Head Typeset by SPi, Pondicherry, India. Printed and bound in the UK by MPG Books Group. Contents Preface to the Second Edition vii 1 The Cat: Domestication and Biology 1 2 Sensory Abilities 16 3 Mechanisms of Behaviour 41 4 Behavioural Development 63 5 Communication 91 6 Feeding Behaviour 113 7 Hunting and Predation 128 8 Social Behaviour 142 9 The Cat–Human Relationship 161 10 Cat Welfare 175 11 Undesired Behaviour in the Domestic Cat 190 12 Physiological and Pathological Causes of Behavioural Change 206 References 217 Index 239 v This page intentionally left blank Preface to the Second Edition The behaviour of the domestic cat, Felis silvestris catus, has many features that set it apart from other common domestic animals, even its fellow carnivore the dog. Cats seem to have effected a unique and successful compromise between reliance on man and the retention of behaviour patterns that permit an independent existence. During the first decade of the 21st century the cat has become the preferred pet of many owners, valued because it combines an affectionate nature with a degree of independence, as well as more prosaic qualities such as cleanliness and convenience. This book brings together many disparate studies of the behaviour of F. catus, and attempts to marry together the more mechanistic approach (how do patterns of behaviour come about?) with the functional (what are those pat- terns for, and how do they benefit the animal?). It may seem unusual to ask functional questions about the behaviour of a domestic animal, but although for many individual cats both feeding and breeding are largely controlled by man, many others live much more independently and are not only subject to natural selection but, through their predatory behaviour, are themselves a selection pressure on other species. This book therefore covers cats across all their current lifestyles, from neutered pet to fully independent (‘feral’). Unlike the domestic dog, which has seen a resurgence in research inter- est over the past decade, several aspects of the cat’s behaviour have received relatively little attention since the publication of the first edition of this book in 1992. These include the cat’s sensory and learning abilities and its repertoire of visual signals. However, major advances have been made in several areas, including cat welfare, social behaviour and domestication, and especially in our understanding and treatment of behavioural disorders, coverage of which has been expanded from one chapter to two. vii viii Preface From a biological perspective, it is often useful to compare one s pecies’ behaviour with that of related species, in this case doubly so, since hybrids between the domestic cat and other small felids are beginning to appear as pets. The first chapter deals with these issues, including the genetic relationship between the domestic cat and the rest of its family, the Felidae, and the basis for its domestication. The next two chapters attempt to describe the world in the cat’s own terms: what it can see, hear, smell and feel, and how it can bring all this incoming information together so that its surroundings make sense. This is worth establishing at the outset, because their perception of the world, while overlapping with ours, also provides them with information that is not directly accessible to us, especially that provided by the cat’s olfactory appa- ratus. Furthermore, cat intelligence has been portrayed in many ways, from the early behaviourists’ picture of a near-automaton to the anthropomorphic sentimentalization of the popular literature; Chapter 3 redresses the balance in favour of modern scientific thinking. Subsequent chapters deal with more spe- cialized aspects of cat behaviour, such as reproduction, development, communi- cation, hunting and feeding. Chapters 8, 9 and 10 are based around a common theme, that of the cat’s social abilities. The complex social interactions between cats, many of which we do not yet fully understand, have forced biologists to reinterpret not only the cat’s relationship with its conspecifics but also the bond with its owner, and considerations of its welfare. The concluding chapters, building on those that precede them, lay out our current understanding of ‘behaviour problems’ in cats, divided into those which are essentially normal behaviours for cats but which owners find undesirable (Chapter 11) and those caused or influenced by pathological or physiological factors (Chapter 12). This framework has been put together primarily by one of us (RAC) from her clinical experience at the University of Bristol’s Animal Behaviour Clinic (recognized as a centre for clinical training by the European College of Animal Welfare and Behavioural Medicine). For those interested in reading further, two recent books can be regarded as complementary to this one. Wild Cats of the World, by Mel and Fiona Sunquist, extends the story to the other members of the Felidae. More com- prehensive treatments of many of the topics are contained in the multi-author book edited by Dennis Turner and Patrick Bateson, The Domestic Cat: the Biology of its Behaviour, which is about to appear in its third edition. Further suggestions can be gleaned from the list of references, which has been deliberately kept short, by using review articles rather than research papers wherever possible, to avoid breaking up the text with large numbers of citations. Many of our friends and colleagues have helped us with this book, by making suggestions, providing references, allowing us to present their unpub- lished research and reading drafts of chapters. In no particular order, they are Sandra McCune, Carlos Driscoll, David Macdonald, Michael Mendl, Ian Robinson, Stuart Church, John Allen, Sarah Lowe (Benge), Ruud van den Bos, Charlotte Cameron-Beaumont, Giles Horsfield, Sarah Hall, Sylvia Vandenbussche, Kim Hawkins, Elizabeth Paul, Carri Westgarth, Emily Preface ix Blackwell, Kristen Kly, Maryanne Heard, Jenna Kiddie, Anne Seawright and Christine Basse. The responsibility for what appears on the printed pages is, of course, ours alone. Our sincere gratitude also goes to Alan Peters for his ink-and-wash sketches of kittens and cats. Our own research, and that of our students, reported here has been supported by the Waltham Centre for Pet Nutrition, the Biotechnology and Biological Sciences Research Council and the Universities Federation for Animal Welfare. We are especially grateful to Cats Protection for supporting research and academic posts over the past two dec- ades, a contribution that has significantly enhanced scientific understanding of cat behaviour and welfare, and its application in clinical practice.