The Domestic Cat. The Biology of its Behaviour. Third Edition

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The Domestic Cat The Biology of its Behaviour Third Edition The most commonly kept domestic animal in the developed world, the cat has been a part of human life for thousands of years. Cats have been both worshipped and persecuted over this long period either loved or hated for their enigmatic self-reliance and the subject of numerous myths and fables. Highlighting startling discoveries made over the last 10 years, this new edition features contributions from experts in a wide range of fields, providing authoritative accounts of the behaviour of cats and how they interact with people. Thoroughly revised and updated to include information on the basic features of cat development and social life, the history of their relations with humans, health and welfare problems, and the breeding of cats for sale and for show, it is intended for all those, whether specialist or general reader, who love or are simply intrigued by these fascinating animals. Dennis C. Turner is Director of the Institute for applied Ethology and Animal Psychology, Switzerland, a private faculty member of the University of Zurich (Vetsuisse Faculty) and Invited Professor at the Educational Research Center for Anthrozoology (ERCAZ), Azabu University, Japan. He has published extensively on domestic cats and is well known for his activities in international organisations concerning human animal relationships, animal-assisted therapy and animal welfare. Patrick Bateson FRS is Emeritus Professor of Ethology at the University of Cambridge. He is President of the Zoological Society of London and former Vice-President of the Royal Society. He is also co-author of Play, Playfulness, Creativity and Innovation (Cambridge, 2013), Plasticity, Robustness, Development and Evolution (Cambridge, 2011) and Measuring Behaviour (Cambridge, 3rd edition 2007).

The Domestic Cat The Biology of its Behaviour Third Edition Edited by DENNIS C. TURNER ScD, Institute for Applied Ethology and Animal Psychology, Hirzel, and PD, Vetsuisse Faculty, University of Zurich, Switzerland PATRICK BATESON FRS, Emeritus Professor of Ethology, Sub-Department of Animal Behaviour, Department of Zoology, University of Cambridge, United Kingdom Ink and wash illustrations by MICHAEL EDWARDS

University Printing House, Cambridge CB2 8BS, United Kingdom Cambridge University Press is part of the University of Cambridge. It furthers the University s mission by disseminating knowledge in the pursuit of education, learning and research at the highest international levels of excellence. Information on this title: /9781107025028 Cambridge University Press 1988, 2014 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 1988 Second edition 2000 Third edition 2014 3rd printing 2015 Printed in the United Kingdom by Clays, St Ives plc. A catalogue record for this publication is available from the British Library Library of Congress Cataloguing in Publication data The domestic cat: the biology of its behaviour / edited by Dennis C. Turner, ScD, Institute for Applied Ethology and Animal Psychology, Hirzel, and PD, Vetsuisse Faculty, University of Zurich, Switzerland, Patrick Bateson, FRS, Emeritus Professor of Ethology, Sub-Department of Animal Behaviour, Department of Zoology, University of Cambridge, United Kingdom; ink and wash illustrations by Michael Edwards. Third edition. pages cm Includes bibliographical references and index. ISBN 978-1-107-02502-8 (Paperback) 1. Cats Behaviour. 2. Cats Social aspects. I. Turner, Dennis C., 1948 II. Bateson, P. P. G. (Paul Patrick Gordon), 1938 SF446.5.D65 2014 636.8 dc23 2013021440 ISBN 978-1-107-02502-8 Paperback Additional resources for this publication at /9781107025028 Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

Contents List of contributors List of abbreviations page vii x I Introduction 1 Why the cat? 3 Dennis C. Turner and Patrick Bateson II From Kitten- to Adulthood 2 Behavioural development in the cat 11 Patrick Bateson 3 Normal and problematic reproductive behaviour in the domestic cat 27 Benjamin L. Hart and Lynette A. Hart 4 Communication in the domestic cat: within- and between-species 37 Sarah L. Brown and John W. S. Bradshaw III Social Life and Ecology 5 Social organisation and behavioural ecology of free-ranging domestic cats 63 Dennis C. Turner 6 Social behaviour of domestic cats in the human home 71 Penny L. Bernstein and Erika Friedmann IV Cats and People 7 Domestication and history of the cat 83 James A. Serpell 8 Cultural differences in human cat relations 101 Dennis C. Turner, Eva Waiblinger and Barbara Fehlbaum

vi Contents 9 Human and cat personalities: building the bond from both sides 113 Kurt Kotrschal, Jon Day, Sandra McCune and Manuela Wedl V Cat Breeding and Cat Welfare 10 Feline welfare issues 131 Irene Rochlitz 11 Breed and gender behaviour differences: relation to the ancient history and origin of the domestic cat 155 Benjamin L. Hart, Lynette A. Hart and Leslie A. Lyons 12 Showing cats 167 Anne Gregory, Steve Crow and Hilary Dean 13 Individual and environmental effects on health and welfare 185 J. L. Stella and C. A. T. Buffington 14 Feline behavioural problems and solutions 201 Benjamin L. Hart and Lynette A. Hart VI The Future 15 Cat population management 215 Elly Hiby, Harry Eckman and Ian MacFarlaine 16 Postscript: questions and some answers 231 Patrick Bateson and Dennis C. Turner References 241 Index 272

Contributors Patrick Bateson Sub-Department of Animal Behaviour, Department of Zoology, University of Cambridge, High Street, Madingley, Cambridge CB23 8AA, UK The Late Penny L. Bernstein formerly Biological Sciences, Kent State University Stark Campus, N. Canton, USA John W. S. Bradshaw School of Veterinary Sciences, University of Bristol, Langford, Bristol BS40 5DU, UK Sarah L. Brown School of Veterinary Sciences, University of Bristol, Langford, Bristol BS40 5DU, UK C. A. T. Buffington Department of Veterinary Clinical Sciences, The Ohio State University College of Veterinary Medicine, Columbus, OH 43210, USA Steve Crow The Governing Council of the Cat Fancy, 5 King s Castle Business Park, Bridgwater, Somerset TA6 4AG, UK Jon Day WALTHAM Centre for Pet Nutrition, Freeby Lane, Waltham-on-the-Wolds, Melton Mowbray, Leicestershire LE14 4RT, UK Hilary Dean The Governing Council of the Cat Fancy, 5 King s Castle Business Park, Bridgwater, Somerset TA6 4AG, UK Harry Eckman Change For Animals Foundation (CFAF), 11 Elmwood Avenue, Borehamwood, Hertfordshire WD6 1SY, UK

viii List of contributors Barbara Fehlbaum V.I.E.T.A., c/o I.E.A.P./I.E.T., PO Box 32, CH-8816 Hirzel, Switzerland Erika Friedmann University of Maryland School of Nursing, Baltimore, MD 21201, USA Anne Gregory The Governing Council of the Cat Fancy, 5 King s Castle Business Park, Bridgwater, Somerset TA6 4AG, UK Benjamin L. Hart School of Veterinary Medicine, Department of Anatomy, Physiology and Cell Biology, University of California-Davis, Davis, CA 95616, USA Lynette A. Hart School of Veterinary Medicine, Department of Population Health and Reproduction, University of California-Davis, Davis, CA 95616, USA Elly Hiby Independent Consultant, 149 Ross Street, Cambridge, CB1 3BS, UK Kurt Kotrschal University of Vienna Department of Behavioural Biology and Konrad Lorenz Forschungsstelle, Althanstraße 14, 1090 Vienna, Austria Leslie A. Lyons School of Veterinary Medicine, Department of Population Health and Reproduction, University of California-Davis, Davis, CA 95616, USA Ian MacFarlaine Head of Operations (Welfare), International Cat Care (formerly Feline Advisory Bureau), Taeselbury, High Street, Tisbury, Wiltshire SP3 6LD, UK Sandra McCune WALTHAM Centre for Pet Nutrition, Freeby Lane, Waltham-on-the-Wolds, Melton Mowbray, Leicestershire LE14 4RT, UK Irene Rochlitz Department of Veterinary Medicine, University of Cambridge, Madingley Road, Cambridge CB3 0ES, UK James A. Serpell Center for the Interaction of Animals & Society (CIAS), School of Veterinary Medicine, University of Pennsylvania, Philadelphia, PA 19104 6010, USA

List of contributors ix J. L. Stella Department of Veterinary Preventive Medicine, The Ohio State University College of Veterinary Medicine, Columbus, OH 43112, USA Dennis C. Turner Institute for Applied Ethology and Animal Psychology, I.E.A.P./I.E.T., PO Box 32, CH-8816 Hirzel, and University of Zurich, Switzerland Eva Waiblinger Head of Companion Animals, Swiss Animal Protection, SAP, Dornacherstrasse 101, PO Box 461, CH-4008 Basel, Switzerland Manuela Wedl University of Vienna Department of Behavioural Biology, Althanstraße 14, 1090 Vienna, Austria

Abbreviations AAFP ACC&D ACTH ADCH ASPCA BCE CAWC CNR CNS CRF DEFRA DJD DLH DSH EE HPA LUTS MYA NCPPSP NRC PCA RSPCA SRS TNR TU VNO WSPA American Association of Feline Practitioners Alliance for Contraception in Cats and Dogs adrenocorticotropic hormone Association of Dogs and Cats Homes American Society for the Prevention of Cruelty to Animals Before the Common Era Companion Animal Welfare Council capture neuter release central nervous system corticotrophin-releasing factor Department for Environment, Food and Rural Affairs degenerative joint disease domestic longhair domestic shorthair environmental enrichment hypothalamus pituitary adrenal lower urinary tract signs Million Years Ago National Council on Pet Population Study and Policy National Research Council Principal Component Analysis Royal Society for the Prevention of Cruelty to Animals stress response system trap neuter return tail up vomeronasal organs World Society for the Protection of Animals