THE BEHAVIOURAL BIOLOGY OF DOGS This page intentionally left blank The Behavioural Biology of Dogs Edited by Per Jensen IFM Biology Linköping University Sweden CABI is a trading name of CAB International CABI Head Office CABI North American Office Nosworthy Way 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org © CAB International 2007. 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. A catalogue record for this book is available from the Library of Congress, Washington, DC. ISBN-10: 1 84593 1874 ISBN-13: 978 1 84593 1872 Typeset by MRM Graphics Ltd, Winslow, Bucks. Printed and bound in the UK by Cromwell Press, Trowbridge. Contents Contributors vii Preface ix I: The Dog in Its Zoological Context 1 Editor’s Introduction 1. Evolutionary History of Canids 3 X. Wang and R.H. Tedford 2. Domestication of Dogs 21 P. Savolainen 3. Origin of Dog Breed Diversity 38 C. Vilà and J.A. Leonard II: Biology and Behaviour of Dogs 59 Editor’s Introduction 4. Mechanisms and Function in Dog Behaviour 61 P. Jensen 5. Behaviour Genetics in Canids 76 E. Jazin v vi Contents 6. Sensory Physiology and Dog Behaviour 91 H. Bubna-Littitz 7. Social Behaviour of Dogs and Related Canids 105 D.U. Feddersen-Petersen 8. Learning in Dogs 120 P. Reid III: The Dog in Its Niche: Among Humans 145 Editor’s Introduction 9. Behaviour and Social Ecology of Free-ranging Dogs 147 L. Boitani, P. Ciucci and A. Ortolani 10. Evolutionary Aspects on Breeding of Working Dogs 166 R. Beilharz 11. Individual Differences in Behaviour – Dog Personality 182 K. Svartberg 12. Human–Animal Interactions and Social Cognition in Dogs 207 Á. Miklósi IV: Behavioural Problems of Dogs 223 Editor’s Introduction 13. Behavioural Disorders of Dogs 225 R.A. Mugford 14. Behaviour and Disease in Dogs 243 Å. Hedhammar and K. Hultin-Jäderlund Index 263 Contributors Beilharz, R., School of Agriculture and Food Systems, Faculty of Land and Food Resources, The University of Melbourne, Melbourne, Victoria 3010, Australia Boitani, L., Department of Animal and Human Biology, University of Rome, Viale Università 32, 00185 Roma, Italy Bubna-Littitz, H., Department for Natural Sciences, Institute for Physiology, Veterinary University of Vienna, Veterinärplatz 1, A-1030 Vienna, Austria Ciucci, P., Department of Animal and Human Biology, University of Rome, Viale Università 32, 00185 Roma, Italy Feddersen-Petersen, D.U., Department of Zoology, University of Kiel, Olshausenstr. 40, D-24118 Kiel, Germany Hedhammar, Å., Department of Small Animal Clinical Sciences, Swedish University of Agricultural Sciences, Box 7037, SE-750 07 Uppsala, Sweden Hultin-Jäderlund, K., Department of Small Animal Clinical Sciences, Norwegian College of Veterinary Medicine, PO Box 8146 Dep., N-0033 Oslo, Norway Jazin, E., Department of Evolution, Genomics and Systematics, Uppsala University, Norbyvagen 18D, SE-752 36 Uppsala, Sweden Jensen, P., IFM Biology, Linköping University, SE-581 83 Linköping, Sweden Leonard, J.A., Department of Evolutionary Biology, Uppsala University, Norbyvägen 18D, SE-752 36 Uppsala, Sweden Miklósi, Á., Department of Ethology, Eötvös Loránd University, Pázmány sétány 1/c 1117, Budapest, Hungary Mugford, R.A., The Animal Behaviour Centre, PO Box 23, Chertsey, Surrey KT16 9NL, UK Ortolani, A., Department of Animal and Human Biology, University of Rome, Viale Università 32, 00185 Roma, Italy Reid, P., ASPCA Animal Behavior Center, 424 East 92nd Street, New York, NY 10128, USA vii viii Contributors Savolainen, P., Department of Biotechnology, Albanova University Centre, KTH-Royal Institute of Technology, SE-10691 Stockholm, Sweden Svartberg, K., Department of Anatomy and Physiology, Swedish University of Agricultural Sciences, PO Box 7011, SE-750 07 Uppsala, Sweden Tedford, R.H., Department of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA Vilà, C., Department of Evolutionary Biology, Uppsala University, Norbyvägen 18D, SE- 752 36 Uppsala, Sweden Wang, X., Department of Vertebrate Paleontology, Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, CA 90007, USA and Department of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA Preface Dogs are our oldest domesticated animals, our friends and companions, and the most widespread species of all animals under human care. There are few things for which dogs have not been utilized during our joint history: they have been used as hunting aids, for pulling and carrying, as providers of meat, fur and other products, for guarding and watching, and as laboratory test animals, just to mention some examples. In large parts of the world, dogs are increasingly popular as pets, friends and family members, and at the same time the importance of dogs as working dogs, for example with police and rescue forces, has probably never been higher. At the same time, biologists have started to take greater interest in the biology and behaviour of dogs, for a number of reasons. First, since dogs are so closely connected to human evolution and history, understanding how, where and when dogs have developed and spread helps us understand our own background. Second, the long coexistence with humans has led the dog to develop specific adaptations facilitating life with us, and the dog therefore provides an excellent model for studying how behaviour and cognition have evolved. Third, the increas- ing importance of dogs as pets and working animals calls for a deeper biological knowledge of how these animals actually work – such information can help us not only to train and shape dogs for specific tasks, but also to prevent and cure various behavioural disorders which may cause owners and animals large problems. The present book is an attempt to provide an up-to-date description of the behavioural biology of dogs, written by experts in different areas of this large field. The target audience consists of students of animal behaviour or veterinary medi- cine at advanced levels – the book is not intended as an introductory text to dog behaviour in general. It is also the hope of the authors that interested dog owners outside academia may find usable parts in the book. There is no doubt that ix x Preface certain chapters will require closer acquaintance with various aspects of biology than the average dog owner is likely to possess, but we also believe that most chapters contain aspects that are readily accessible. The book is split into four different parts, each concerned with a specific aspect of the behavioural biology of dogs. The first part (Chapters 1–3) is devoted mainly to the evolution and development of the dog. Although not primarily con- cerned with behaviour, these aspects form the basis for understanding how behaviour has developed and for placing the dog in its relevant biological context. The second part (Chapters 4–8) deals with basic aspects of animal behaviour with particular emphasis on dogs. The third part (Chapters 9–12) places the modern dog in its present ecological framework: in the niche of human coexistence. Here we give a broad overview of the behavioural aspects of living close to humans. In the last part of the book (Chapters 13 and 14), the emphasis is on behavioural problems, their prevention and cure. All of the contributors to this book have considerable research experience in their areas, and it is hoped that this will guarantee that the text is relevant, up-to- date and central to the subject. Per Jensen Linköping, March 2006 I The Dog in Its Zoological Context Editor’s Introduction In this first part of the book, the dog is placed in context amongst its zoological relatives, and in relation to its domestication history. The first chapter outlines the modern view on the zoological systematics of carnivores and canids in particular. Here, the reader will find an exciting account of fossil and present traits which allows the dog to be placed within the greater picture of closely related canids. Of course, the dog is a domesticated species, and its domestication history has been subject to intense research during the last decade or so. Here, modern molecular genetics offers tools which have allowed biologists to give their pictures of how, where and when domestication started, complementing the traditional picture offered mainly by archaeologists. Some aspects of this new picture are truly stunning and require that we revise large parts of our traditional views on how domestication began. Given the novelty of the molecular research on the ancestry of dogs, it should not be a surprise that different scientific groups arrive at somewhat different con- clusions. This is partly explained by variations in methods, and the only way to resolve some of the disagreements is to continue to do more and improved research. As this book is written, there is therefore only limited consensus among biologists concerning, for example, the time when domestication started. As editor, one has to make a decision – one can choose not to cover the new research at all, or to only present the picture oneself believes in, or to present divergent pic- tures and allow the reader to decide. I have chosen the last strategy, and there- fore, the stories presented in Chapters 2 and 3 differ on some important points – these differences will hopefully disappear in the light of future research.