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Status Survey and Conservation Action Plan Canids: Foxes, Wolves, Jackals and Dogs Edited by Claudio Sillero-Zubiri, Michael Hoffmann and David W. Macdonald IUCN/SSC Canid Specialist Group

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  • Canids: Foxes, W

    olves, Jackals and Dogs

    Statu

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    IUCN/Species Survival Commission

    The Species Survival Commission (SSC) is one of six volunteer commissions of IUCN – TheWorld Conservation Union, a union of sovereign states, government agencies and non-governmental organisations. IUCN has three basic conservation objectives: to secure theconservation of nature, and especially of biological diversity, as an essential foundation for thefuture; to ensure that where the Earth’s natural resources are used this is done in a wise,equitable and sustainable way; and to guide the development of human communities towardsways of life that are both of good quality and in enduring harmony with other components of thebiosphere.

    A volunteer network comprised of some 8,000 scientists, field researchers, government officialsand conservation leaders from nearly every country of the world, the SSC membership is anunmatched source of information about biological diversity and its conservation. As such, SSCmembers provide technical and scientific counsel for conservation projects throughout theworld and serve as resources to governments, international conventions and conservationorganisations.

    IUCN/SSC Action Plans assess the conservation status of species and their habitats, andspecifies conservation priorities. The series is one of the world’s most authoritative sources ofspecies conservation information available to natural resource managers, conservationists,and government officials around the world.

    IUCN Species ProgrammeRue Mauverney 28, CH-1196 Gland, SwitzerlandTel: +41 22 999 00 00, Fax: +41 22 999 00 15E-mail: [email protected]/themes/ssc

    IUCN Publications Services Unit219c Huntingdon Road, Cambridge, CB3 0DL, UKTel: +44 1223 277894, Fax: +44 1223 277175E-mail: [email protected]/bookstore

    Status Survey and Conservation Action Plan

    Canids: Foxes, Wolves,Jackals and Dogs

    Edited by Claudio Sillero-Zubiri, Michael Hoffmannand David W. Macdonald

    IUCN/SSC Canid Specialist Group

  • Donors to the SSC Conservation Communications Programme andCanids: Foxes, Wolves, Jackals and Dogs Status Survey and Conservation Action Plan

    The IUCN Species Survival Commission is committed to communicate important species conservation information tonatural resource managers, decision makers and others whose actions affect the conservation of biodiversity. The SSC’sAction Plans, Occasional Papers, newsletter Species and other publications are supported by a wide variety of generousdonors including:

    The Sultanate of Oman established the Peter Scott IUCN/SSC Action Plan Fund in 1990. The Fund supports Action Plandevelopment and implementation. To date, more than 80 grants have been made from the Fund to SSC Specialist Groups.The SSC is grateful to the Sultanate of Oman for its confidence in and support for species conservation worldwide.

    The Council of Agriculture (COA), Taiwan has awarded major grants to the SSC’s Wildlife Trade Programme andConservation Communications Programme. This support has enabled SSC to continue its valuable technical advisoryservice to the Parties to CITES as well as to the larger global conservation community. Among other responsibilities, theCOA is in charge of matters concerning the designation and management of nature reserves, conservation of wildlife andtheir habitats, conservation of natural landscapes, coordination of law enforcement efforts, as well as promotion ofconservation education, research and international cooperation.

    The World Wide Fund for Nature (WWF) provides significant annual operating support to the SSC. WWF’s contributionsupports the SSC’s minimal infrastructure and helps ensure that the voluntary network and publications programme areadequately supported. WWF aims to conserve nature and ecological processes by: (1) preserving genetic, species, andecosystem diversity; (2) ensuring that the use of renewable natural resources is sustainable both now and in the longerterm; and (3) promoting actions to reduce pollution and the wasteful exploitation and consumption of resources andenergy. WWF is one of the world’s largest independent conservation organisations with a network of NationalOrganisations and Associates around the world and over 5.2 million regular supporters. WWF continues to be knownas World Wildlife Fund in Canada and in the United States of America.

    Donors to the Canid Specialist Group and Canids: Foxes, Wolves, Jackals and DogsThe Canid Specialist Group is the world’s chief body of scientific and practical expertise on the status and conservationof all species of the Canidae, advising the IUCN Species Survival Commission. The CSG has its base at the WildlifeConservation Research Unit (WildCRU) and is composed of more than 100 experts, representing over 30 countries (andwith experience in many more), serving as honorary advisors.

    The Wildlife Conservation Research Unit of the University of Oxford (known as the WildCRU) seeks to achieve practicalsolutions to conservation problems. WildCRU does this through original scientific research of the highest calibre. Vitally,it also trains committed conservation scientists to conduct research, and to put scientific knowledge into practice.WildCRU participates in the implementation of many conservation projects and embraces the need to educate andinvolve a wider public to achieve lasting solutions.

    The Born Free Foundation is an international wildlife charity, founded by actors Virginia McKenna and Bill Traversfollowing their starring roles in the classic film ‘Born Free’. Born Free works to alleviate suffering, investigate cruelty andencourage everyone to treat wildlife with respect. Born Free believes wildlife belongs in the wild and is dedicated to theprotection of threatened species in their natural habitat, working with local communities around the world to help peopleand wildlife live together without conflict.

    The People’s Trust for Endangered Species is committed to working to preserve wildlife in their natural habitats for futuregenerations to enjoy. To achieve this aim PTES commissions and funds research, carries out practical work in the field,purchases reserves, runs an education programme, provides opportunities for members of the public to take part innationwide surveys and organises national and international conferences.

  • Status Survey and Conservation Action Plan

    Canids: Foxes, Wolves,Jackals and Dogs

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  • iii

    Status Survey and Conservation Action Plan

    Canids: Foxes, Wolves,Jackals and Dogs

    Edited by Claudio Sillero-Zubiri, Michael Hoffmannand David W. Macdonald

    IUCN/SSC Canid Specialist Group

    IUCN – The World Conservation Union2004

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    The designation of geographical entities in this book, and the presentation of the material, do not imply the expression of any opinionwhatsoever on the part of IUCN concerning the legal status of any country, territory, or area, or of its authorities, or concerning thedelimitation of its frontiers or boundaries.

    The views expressed in this publication do not necessarily reflect those of IUCN.

    Published by: IUCN, Gland, Switzerland and Cambridge, UK

    Copyright: © 2004 International Union for Conservation of Nature and Natural Resources

    Reproduction of this publication for educational or other non-commercial purposes is authorised without prior writtenpermission from the copyright holder provided the source is fully acknowledged.

    Reproduction of this publication for resale or other commercial purposes is prohibited without prior written permission ofthe copyright holder.

    Citation: Sillero-Zubiri, C., Hoffmann, M. and Macdonald, D.W. (eds). 2004. Canids: Foxes, Wolves, Jackals and Dogs. Status Surveyand Conservation Action Plan. IUCN/SSC Canid Specialist Group. Gland, Switzerland and Cambridge, UK. x + 430 pp.

    ISBN: 2-8317-0786-2

    Cover photo: Male Blanford’s fox (Vulpes cana). Many fox species can be easily identified by large ears that have developed to enablethe animals to pick up the minute rustlings of small prey, and to help dissipate heat in hot climates. Jebel Hafit, Al Ain, AbuDhabi, United Arab Emirates, 1998. Photo: Chris and Tilde Stuart.

    Produced by: The Wildlife Conservation Research Unit, Oxford, UK

    Photo editor: Michelle Nelson

    Proofreader: Sue Sefscik, Polly Phillpot and Bryan Hugill

    Mapsprepared by:

    WildCRU in collaboration with Wes Sechrest and the Global Mammal Assessment

    Layout by: The NatureBureau, Newbury, UK

    Printed by: Information Press, Oxford, UK

    Available from: IUCN Publications Services Unit219c Huntingdon Road, Cambridge CB3 0DL, United KingdomTel: +44 1223 277894, Fax: +44 1223 277175E-mail: [email protected]/bookstoreA catalogue of IUCN publications is also available.

    The text of this book is printed on 90 gsm Fineblade Smooth, which is rmade from 100% sustainable fibre sources using chlorine-freeprocesses.

  • v

    Contents

    Foreword .....................................................................vii

    Acknowledgements .....................................................viii

    Executive Summary ....................................................... x

    PART 1Overview ....................................................................... 1

    Chapter 1. Introduction ................................................. 2CLAUDIO SILLERO-ZUBIRI AND DAVID W. MACDONALD

    Chapter 2. Phylogeny, Classification, andEvolutionary Ecology of the Canidae ............................. 8WANG XIAOMING, RICHARD H. TEDFORD,BLAIRE VAN VALKENBURGH AND ROBERT K. WAYNE

    PART 2Species Status Accounts .............................................. 21EDITED BY MICHAEL HOFFMANN AND CLAUDIO SILLERO-ZUBIRI

    Species Status Accounts: an Introduction .................... 22

    Chapter 3. South America (Neotropical) ...................... 263.1 Short-eared dog (Atelocynus microtis) ................. 26

    M. RENATA P. LEITE PITMAN AND ROBERT S.R. WILLIAMS

    3.2 Crab-eating fox (Cerdocyon thous) ...................... 32ORIN COURTENAY AND LEONARDO MAFFEI

    3.3 Maned wolf (Chrysocyon brachyurus) ................. 38MELISSA RODDEN, FLÁVIO RODRIGUESAND STEPHANIE BESTELMEYER

    3.4 Culpeo (Pseudalopex culpaeus) ............................ 44JAIME E. JIMÉNEZ AND ANDRÉS J. NOVARO

    3.5 Darwin’s fox (Pseudalopex fulvipes) .................... 50JAIME E. JIMÉNEZ AND ELISE MCMAHON

    3.6 Chilla (Pseudalopex griseus) ................................ 56RAFAEL GONZÁLEZ DEL SOLAR AND JAIME RAU

    3.7 Pampas fox (Pseudalopex gymnocercus) .............. 63MAURO LUCHERINI, MARCELO PESSINOAND ARIEL A. FARIAS

    3.8 Sechuran fox (Pseudalopex sechurae) .................. 69CHERYL ASA AND E. DANIEL COSSÍOS

    3.9 Hoary fox (Pseudalopex vetulus) ......................... 72JULIO DALPONTE AND ORIN COURTENAY

    3.10 Bush dog (Speothos venaticus) ............................. 76GERALD L. ZUERCHER, MATTHEW SWARNER,LEANDRO SILVEIRA AND OSVALDO CARRILLO

    Chapter 4. Central and North America (Nearctic) ....... 814.1 Coyote (Canis latrans) ......................................... 81

    ERIC M. GESE AND MARC BEKOFF4.2 Red wolf (Canis rufus) ......................................... 87

    BRIAN T. KELLY, ARTHUR BEYERAND MICHAEL K. PHILLIPS

    4.3 Gray fox (Urocyon cinereoargenteus) .................. 92TODD K. FULLER AND BRIAN L. CYPHER

    4.4 Island fox (Urocyon littoralis) ............................. 97GARY W. ROEMER, TIMOTHY J. COONAN,LINDA MUNSON AND ROBERT K. WAYNE

    4.5 Kit fox (Vulpes macrotis) ................................... 105RURIK LIST AND BRIAN L. CYPHER

    4.6 Swift fox (Vulpes velox) ..................................... 109AXEL MOEHRENSCHLAGER AND MARSHA SOVADA

    Chapter 5. Europe and North and Central Asia(Palearctic) ................................................................ 1175.1 Arctic fox (Alopex lagopus) ............................... 117

    ANDERS ANGERBJÖRN, PALL HERSTEINSSON ANDMAGNUS TANNERFELDT

    5.2 Grey wolf (Canis lupus) ..................................... 124L. DAVID MECH AND LUIGI BOITANI

    5.3 Red fox (Vulpes vulpes) ...................................... 129DAVID W. MACDONALD AND JONATHAN REYNOLDS

    5.4 Raccoon dog (Nyctereutes procyonoides) .......... 136KAARINA KAUHALA AND MIDORI SAEKI

    5.5 Corsac (Vulpes corsac) ....................................... 142ANDREI POYARKOV AND NIKITA OVSYANIKOV

    5.6 Tibetan fox (Vulpes ferrilata) ............................ 148GEORGE B. SCHALLER AND JOSHUA R. GINSBERG

    Chapter 6. Sub-Saharan Africa (Ethiopian) ............... 1526.1 Side-striped jackal (Canis adustus) .................... 152

    ROBERT P.D. ATKINSON AND ANDREW J. LOVERIDGE

    6.2 Golden jackal (Canis aureus) ............................. 156YADVENDRADEV V. JHALA ANDPATRICIA D. MOEHLMAN

    6.3 Black-backed jackal (Canis mesomelas) ............ 161ANDREW J. LOVERIDGE AND JAN A.J. NEL

    6.4 Ethiopian wolf (Canis simensis) ......................... 167CLAUDIO SILLERO-ZUBIRI AND JORGELINA MARINO

    6.5 African wild dog (Lycaon pictus) ....................... 174ROSIE WOODROFFE, J. WELDON MCNUTTAND M.G.L. MILLS

    6.6 Bat-eared fox (Otocyon megalotis) .................... 183JAN A.J. NEL AND BARBARA MAAS

    6.7 Cape fox (Vulpes chama) ................................... 189CHRIS STUART AND TILDE STUART

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    Chapter 7. North Africa andthe Middle East (Ethiopian) ....................................... 1947.1 Blanford’s fox (Vulpes cana) .............................. 194

    ELI GEFFEN, REUVEN HEFNER AND PETER WRIGHT7.2 Pale fox (Vulpes pallida) .................................... 199

    CLAUDIO SILLERO-ZUBIRI7.3 Rüppell’s fox (Vulpes rueppellii) ........................ 201

    FABRICE CUZIN AND DANNY M. LENAIN7.4 Fennec fox (Vulpes zerda) .................................. 205

    CHERYL S. ASA, CAROLINA VALDESPINOAND FABRICE CUZIN

    Chapter 8. South Asia – South of the Himalaya(Oriental) .................................................................. 2108.1 Dhole (Cuon alpinus) ......................................... 210

    LEON S. DURBIN, ARUN VENKATARAMAN,SIMON HEDGES AND WILL DUCKWORTH

    8.2 Indian fox (Vulpes bengalensis) ......................... 219A.J.T. JOHNSINGH AND YADVENDRADEV V. JHALA

    Chapter 9. Australia and Oceania (Australasian) ...... 2239.1 Dingo (Canis lupus dingo) .................................. 223

    L.K. CORBETT

    PART 3Major issues in Canid Conservation ........................... 231

    Chapter 10. Canid Sociology – a Brief Overview ....... 232DAVID W. MACDONALD

    Chapter 11. Conservation Genetics of Canids ............ 237ROBERT K. WAYNE, ELI GEFFEN AND CARLES VILÀ

    Chapter 12. Assessing and Managing InfectiousDisease Threats to Canids ......................................... 246M. KAREN LAURENSON, SARAH CLEAVELAND,MARC ARTOIS AND ROSIE WOODROFFE

    Chapter 13. Management of Wild Canids inHuman-Dominated Landscapes ................................. 256CLAUDIO SILLERO-ZUBIRI AND DAVID SWITZER

    Chapter 14. Evaluating and Predicting the Impactsof Exploitation and Trade on Canid Populations ....... 267WARREN E. JOHNSON

    Chapter 15. Survey and Census Techniquesfor Canids .................................................................. 273ERIC GESE

    Chapter 16. Captive Canid Conservation ................... 280KAREN L. BAUMANN, CHERYL S. ASA, JACK GRISHAMAND WIM VERBERKMOES

    Chapter 17. Canid Reintroductions andMetapopulation Management .................................... 289AXEL MOEHRENSCHLAGER AND MICHAEL SOMERS

    Chapter 18. Conservation Education andits Relevance to Wild Canids ..................................... 298DENISE TAYLOR

    PART 4Action Plan for Canid Conservationinto the 21st Century ................................................. 307

    Chapter 19. Some Considerations forSetting Priorities and Measuring Successfor Canid Conservation .............................................. 308M. GUS L. MILLS

    Chapter 20. Action Plan for Canid Conservationinto the 21st Century ................................................. 310EDITED BY CLAUDIO SILLERO-ZUBIRI, DAVID W. MACDONALDAND THE CANID AND WOLF SPECIALIST GROUPS

    References ................................................................. 343

    Appendix 1: List of Canids on the2004 IUCN Red List of Threatened Species .............. 404

    Appendix 2: IUCN Red List Categories and CriteriaVersion 3.1 ................................................................ 405

    Appendix 3: List of Contributors Addresses ............... 419

    Appendix 4: List of Reviewers andProject Contacts ........................................................ 423

    Appendix 5: Canid Specialist Group Resources ......... 427

  • vii

    Foreword

    As humans usurp more and more of the Earth, and thenatural world continues to shrink, carnivores will bear adisproportionate toll of the effects. This is becausecarnivores tend to have larger home ranges, more extensivemovements, and longer dispersal distances than theirprey, so their spatial requirements bring them into greatercontact with humans. Furthermore, carnivores tend toconflict directly with human interests because of theproclivity of many of them to kill animals that humans usethemselves.

    Canids form one of the most prominent families ofcarnivores, with 36 interesting taxa in 13 genera that occurthroughout most of the world. Foxes, dholes, dingoes,wolves, jackals, coyotes and various dogs comprise thefamily, and they find human-raised livestock irresistibleprey. As a family, canids occupy every continent exceptAntarctica. The grey wolf, alone, was originally the mostwidely distributed terrestrial mammal; its successor to thethrone is another successful canid, the red fox. Thus,canids have borne a high proportion of the conflict betweenhumans and carnivores. The more prolific and adaptablecanids, like the jackal and coyote, have fared well despitethis competition, while the more specialised members ofthe family, like the Ethiopian wolf, have become threatenedwith extinction.

    However, whatever the past or present status of aparticular canid species, we can be sure that the future willpresent new problems as human populations grow, intrudeon natural habitat, and convert more of the Earth to their

    own liking. Whether the issue is habitat loss, directcompetition, or disease spread (both from canids to humansas with rabies, or from human sources to canids, such ascanine parvovirus from domestic dogs to wolves), increasinghuman pressure means that canids face an uncertain future.

    Fortunately, some humans have taken notice and havedecided to assess the situation systematically. Throughthe World Conservation Union’s (IUCN) Species SurvivalCommission, the Canid Specialist Group has developedthis Action Plan for Canid Conservation. Editors ClaudioSillero-Zubiri, Michael Hoffmann and David Macdonaldhave assembled here an impressive collection ofinformation about all living canid species and theconservation problems they face. From genetics to diseases,conflict resolution to reintroduction, this Canid ActionPlan not only covers the basics, but also addresses themost pressing issues for canid conservation in acomprehensive manner.

    Perusing this wealth of well-organised and importantinformation is enough to give one hope that, despite themany problems canids face, this mobilisation ofinformation and planning will help ensure the survival ofall these intriguing species that comprise the familyCanidae.

    L. Dave MechChair, IUCN/SSC Wolf Specialist GroupU.S. Geological SurveySt. Paul, Minnesota, USA

    Eight-week old island fox pup(Urocyon littoralis). Santa RosaIsland, California, USA, 2001.

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  • viii

    Acknowledgements

    This book has grown from the deliberations of the IUCN/SSC Canid Specialist Group, during many gatheringssince the Group was formed in 1981. This exchange ofideas culminated in September 2001 with the Canid Biologyand Conservation Conference at Oxford, where many ofthe ideas presented in this volume were discussed. We oweour gratitude to each of the 88 authors that contributed tothis volume (Appendix 3), most of whom are members ofthe Canid Specialist Group. The Group is supported by anarmy of affiliates around the world, whose efforts haveproduced the data on which this action plan are based (seeAppendix 4). Our gratitude goes to all that generouslycorresponded with us, reviewed manuscripts or assistedwith data.

    Grey and red wolves are watched over by a separateWolf Specialist Group, and we are grateful to that groupand, in particular, its chairman, Dave Mech, for supplyingthe information we present on grey and red wolves. We arealso grateful to Mariano Gimenez-Dixon, Polly Philpotand Bryan Hugill at the Species Survival Commissionoffice in Gland for reviewing and overseeing the productionof this Action Plan.

    Preparation of the Plan has involved a mountain ofcorrespondence and internet work, the burden of whichfell largely on Claudio’s shoulders. He received much ableassistance from David Switzer, Deborah Randall,Guillaume Chapron and Jorgelina Marino. We wouldparticularly like to thank Sue Scefsik for proof-readingevery section of the manuscript, Deborah Randall whoassisted editing several species’ accounts, and SamBremner-Harrison for additional proof-reading. WillDuckworth provided useful advice on the use of commonEnglish names. Wes Sechrest of the IUCN/SSC – CI/CABS Global Mammal Assessment is thanked forsupplying GMA map templates and Ruaridh Campbell atWildCRU for producing the final maps.

    We are extremely grateful to Michelle Nelson forassembling a wonderful collection of photographs toillustrate this volume. The Canid Specialist Group wouldlike to express its gratitude to the following people thatdonated the use of their photographs, many of which areportrayed in this volume: Lee Allen, Anders Angerbjörn,Daniel Ascensios, Stephanie Bestelmeyer, Art Beyer,Karen Brown, William Campbell, Timothy J. Coonan,Enrique Couve Montané, Barron Crawford, RogerioCunha, Love Dalén, Marcelo Dolsan, Chuck Dresner,Leon Durbin, Eric Gese, Great Tanuki Club, AdrianoGambarini, Rafael González del Solar, Páll Hersteinsson,Don Jones, Steve Kaufman, Greg Koch, Renata LeitePitman, Rurik List, Andrew Loveridge, David Macdonald,Kim McCreery, Elise McMahon, Michal Murri, Michelle

    Nelson, Andrés Novaro, Travis Olson, Hira Punjabi,Robert Robbins, Gary Roemer, Flávio Rodrigues, MidoriSaeki, George Schaller, Krupakar Senani, DavidShackleton, Hólmfríður Sigþórsdóttir, Claudio Sillero,C. Smith, Stanley Park Ecology Society, Chris and TildeStuart, Adam Surgenor, Matt Swarner, MagnusTannerfeldt, Steve Wadlow, Lisa Ware, Wildlife RescueAssociation of British Columbia, and Gerald Zuercher.

    Thanks to the Born Free Foundation and the Peoples’Trust for Endangered Species, which have generouslysupported the Canid Specialist Group and severalWildCRU projects and contributed towards theproduction of this Action Plan. The Zoology Departmentat Oxford University kindly hosted the Canid Biology andConservation Conference in 2001. Thanks to the IUCN/SSC Peter Scott Action Plan Fund that contributed towardsthe cost of this publication and Conservation Internationalfor USA distribution.

    We are particularly grateful to Lady Margaret Hall,the Oxford College where we both hold Fellowships. TheBorn Free Foundation has supported Claudio there whilehe edited this volume. Part of David’s work on this ActionPlan was supported by the Peoples’ Trust for EndangeredSpecies, while some of Mike’s time was supported by theCenter for Applied Biodiversity Science at ConservationInternational. The production of this plan was closelylinked to the Oxford conference, and many of the 250international delegates were accommodated at LMH. Intotal then, this venture owes much to the College, and tothe enthusiastic support of its Principal, which we warmlyacknowledge.

    The authors of Chapter 2 appreciate the review of theirwork by Olaf R.P. Bininda-Emonds.

    The authors of short-eared dog account would liketo thank A. Gardner, J. Patton, H. Beck, A. Forsyth,T. Defler, M. Messias, M. Vivo, C. Canaday, L. Jost,L. Coloma, F. Campos Y., M. Foster, L. Lopez,M. Silman, S. Miller, R. Wallace, F. Wilkinson, A. Salas,D. Wilamowski, M. Boddicker, P. Santos, G. Rosenberg,M. Lelish and S. Ferrari, who shared their fieldobservations; M. Hafner, B. Stanley, B. Patterson,V. Pacheco, J. Amanzo, A. Percequillo, O. Vaccaro,D. Hills, S. Maris, L. Salles, S. Marques-Aguiar,C. Indrusiak and T. Pacheco, who helped with museumdata; A. Shoemaker, D. Kleiman, M. Murphy, C. Arias,for helping with data on captive animals; C. Canaday,S. Athayde and M. Swarner, for providing details ofindigenous names; and N. Pitman, who helped preparethe manuscript. Special thanks go to J. Terborgh, whoencouraged and stimulated this work from its inception;and to Disney Conservation Fund, Conservation

  • ix

    International, IdeaWild, and Wildlife Materials forfinancial support.

    The authors of the crab-eating fox account thank thefollowing for providing abstracts and unpublishedinformation when preparing this review: C. Baltzinger, J.Cartes, F. Catzeflis, J. Chebez, D. Cossios, J. Cravino, R.Cunha de Paula, J. Dalponte, S. Deem, C. Delgado, B.Espinosa, K. Juarez, M. Fabian, K. Facure, A. Farias, J.Tavares Filho, A. Giaretta, R. Leite, B. Lim, M. Lucherini,Y. Martínez, R. Rodrígues-Mazzini, F. Michalski, A.Novaro, L. Pacheco, A. Parera, R. de Paula, R. Printes,D. Queriolo, D. Rode, F. Rodrigues, R. Rodríguez-Mazzini, L. Soler, A. Soutullo, J. Tavares-Filho, R. Varela,R. Williams, and G. Zuercher. Their sincere gratitude alsoto J. Dalponte, R. Leite, A. Parera, and M. Lucherini forhelpful suggestions on earlier versions of the manuscript.

    The authors of the maned wolf account gratefullyacknowledge M. Beccaceci, L. Soler, E. Richard, D. Rumiz,L. Sainz, L. Emmons, R. Wallace, J.R. Moreira, C.B.X.da Silva, F. Michalski, G. Zuercher and S. Gonzalez, all ofwhom provided additional information about populationestimates, distribution, habitat use and diet. J.R. Moreiraassisted in the compilation of the distribution map.

    The authors of the bush dog account thank H. Chaves,P. Crawshaw, J.C. Dalponte, P. Keller-Colcombet, C.Mercolli, T. de Oliveira, M. Swarner and R. Wallace forproviding additional details of protected areas wherebush dogs are reported to occur.

    The authors of the Darwin’s fox account appreciatethe support of the Research Division of Universidad deLos Lagos, IdeaWild, and the Lincoln Park Zoo of Chicagowho have funded J.E. Jiménez’ research on the Darwin´sfox on Chiloé Island. E. McMahon is funded by a GAANNResearch Fellowship from the Graduate Program inOrganismic and Evolutionary Biology and a researchgrant from the Department of Natural ResourcesConservation at the University of Massachusetts, theNational Geographic Society, a grant from the MargaretE. and Howard E. Bigelow Gift to the University ofMassachusetts Amherst, and the Woods Hole ResearchFund. Thanks to Agustin Iriarte and the many other

    people from SAG and CONAF who have supported themainland research of the Darwin’s fox, and all the fieldassistants and park rangers who have helped in this study.

    The authors of Chapter 11 would like to thank K.Koepfli and B. Van Valkenburgh for helpful comments.Their work was supported by the U.S. Fish and WildlifeService, the National Science Foundation, The NatureConservancy, Wolf Haven, the Smithsonian Institution,National Geographic Society, El Padron, and the GeneticResources and Conservation Program and the AcademicSenate of the University of California.

    The author of Chapter 14 would like to thank GregRasmussen for helpful comments on an earlier manuscript.

    Eric Gese (Chapter 15) wishes to thank W. Andelt, A.Angerbjorn, S. Creel, F. Cuzin, K. DeMatteo, J. Dietz, T.Fuller, P. Hersteinsson, K. Karanth, K. Laurenson, M.Lucherini, J. Malcolm, P. Phelan, J. Rau, F. Rodrigues,G. Roemer, C. Sillero, and E. Vidal for providinginformation about methods they have employed on theirprojects.

    Axel Moehrenschlager (Chapter 17) would like tothank Wildlife Preservation Trust Canada and HuskyEnergy for funding support towards this study, LukeHunter for his critical review, and Jay Gedir for editorialcomments.

    Denise Taylor (Chapter 18) would like to thankeveryone who has supported and influenced herconservation education work. In particular, ProfessorLeicester, Professor Susan Jacobson and her friends andcolleagues in wolf conservation that have providedinvaluable support. Guillaume Chapron and ChristophPromberger kindly reviewed an earlier manuscript.

    The production of this action plan has involved muchhard work and generosity by many people andorganisations, and it is hoped that this support will alsofacilitate its implementation.

    David W. Macdonald,Chair IUCN/SSC Canid Specialist GroupClaudio Sillero-Zubiri,Deputy Chair IUCN/SSC Canid Specialist Group

  • x

    Executive Summary

    than half of the mortality recorded among adults is causeddirectly by human activity.

    • Dhole (EN). Formerly distributed across Asia, dholes haveundergone widespread decline and are threatened bydepletion of their prey base, habitat loss, persecution,competition and disease.

    • Dingo (VU). Austronesian people transported the dingofrom Asia to Australia and other islands in between 1,000and 5,000 years ago. Pure dingoes occur only as remnantpopulations in central and northern Australia and inThailand, and they are threatened by cross-breeding withdomestic dogs.

    • Bush dog (VU). Despite a supposedly widespread distributionin South American forests, this species is perceived as rare,and threatened by habitat conversion and humanencroachment.

    • Blanford’s fox (VU). Present in arid mountainous regions ofthe Middle East and north-eastern Egypt eastwards toAfghanistan, where human development could pose a threat.

    In contrast to the threats faced by threatened canids, severalspecies are thriving in human-dominated landscapes and incurthe loathing of farmers and hunters alike. Red foxes arenotoriously successful in urban settings, and coyotes, goldenjackals, crab-eating and kit foxes seem able to thrive amidsthuman settlements. Management prompted by rabies control,fur harvest, and livestock predation leads to the slaughter ofhundreds of thousands of canids annually.

    Part 3 of the Plan considers nine major issues in canidconservation, namely canid society, conservation genetics,assessing and managing diseases, management of canids nearpeople, impact of exploitation and trade, survey and censustechniques, captive conservation, reintroduction and meta-population management, and conservation education.

    Part 4 is arguably the most important section. It includes achapter on the need for setting priorities and measuring successin canid conservation, and the detailed Action Plan for canidconservation into the 21st century. Although we have sought torefine and consolidate these entries, they represent the views ofthe many experts around the world who suggested them, whodebated them in our workshops and in the forum of ourinternational congress, hosted by the WildCRU in Oxford inSeptember 2001. The list of proposed projects makes no claim tobe comprehensive, but it is the result of extremely wideconsultation. The plan itself, together with the databasesconcerning existing members and research projects, are allavailable on the web at http://www.canids.org. The Action Planwas prepared in parallel with our edited monograph entitled TheBiology and Conservation of Wild Canids (Oxford UniversityPress, 2004) which contains comprehensive reviews of the scienceunderpinning this Action Plan, together with 14 case studies ofwild canid biology.

    David W. Macdonald,Chairman IUCN/S SC Canid Specialist GroupClaudio Sillero-Zubiri,Deputy Chair IUCN/S SC Canid Specialist Group

    The new Canid Action Plan synthesises the current knowledgeon the biology, ecology and status of all wild canid species, andoutlines the conservation actions and projects needed to securetheir long-term survival. Aiming at conservation biologists,ecologists, local conservation officials, administrators, educators,and all others dealing with canids in their jobs, the authors aspireto stimulate the conservation of all canids by highlightingproblems, debating priorities and suggesting action.

    The 36 taxa of wild canids that comprise the family Canidae,ranging in size from the tiny fennec fox to the mighty grey wolf,and found in every continent except Antarctica, are special. Theyare special because they have, as perceived friend or foe,preoccupied the imaginations of mankind for millennia; becausethe breadth of their adaptations makes them enthralling toscience; and because the contradictory facets of their relationswith people perplex the conservationist. The increase in numbersof people, the spread of settlement, and the exploitation ofnatural resources of previously little-disturbed wild lands, togetherwith persecution, are threatening some of these canids withextinction. The possibility that we are heedlessly, perhapsneedlessly, mismanaging many of them is saddening; theprobability that our negligence will force several more to extinctionshould fill us with bottomless dismay. It demands action, andthat is why we have compiled this new Canid Action Plan.

    Following a short introduction and a chapter on phylogeny,classification, and evolutionary ecology of the Canidae (Part 1),Part 2 provides the latest information on the distribution, biologyand conservation status of each species, organised by geographicalregion. The accounts also list current field projects, and theircontact details are provided in an appendix. The Canid SpecialistGroup’s members are active worldwide. Nine of the 36 taxacovered are threatened (3 Critically Endangered, 3 Endangeredand 3 Vulnerable), and one is considered Near Threatened. Sixspecies (7%) were listed as Data Deficient, and 20 (56%) speciesas Least Concern (Appendix 1). The threatened canids are:• Darwin’s fox (CR). Until recently, known only from the

    Island of Chiloé (Chile) until rediscovered 600km away inthe coastal mountains, where domestic dogs threaten themwith disease or direct attack.

    • Red wolf (CR). Currently the subject of taxonomic debate,red wolves were declared Extinct in the Wild by 1980, buthave been reintroduced into eastern North Carolina, wherethey are now locally common. Hybridisation with coyotes isthe primary threat.

    • Island fox (CR). Restricted to the six largest of the eightCalifornia Channel Islands, each island population isconsidered a separate subspecies, and four have declinedprecipitously. Threats include hyperpredation by goldeneagles and the introduction of canine diseases.

    • Ethiopian wolf (EN). Less than 500 individuals remain,confined to eight locations in the Ethiopian Highlands.Previously listed as Critically Endangered, continuous lossof habitat due to high-altitude subsistence agriculture remainsthe major threat, along with disease (particularly rabies).

    • African wild dog (EN). Formerly distributed throughoutsub-Saharan Africa, excluding rainforests, wild dogs havedisappeared from 25 of the former 39 range states. More

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    PART 1

    Overview

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    Chapter 1

    IntroductionC. Sillero-Zubiri and D.W. Macdonald

    Canids: Foxes, Wolves, Jackals and Dogs Status Surveyand Conservation Action Plan consists of a review of 36wild canids, of which at least nine are threatened withextinction: island fox (Urocyon littoralis), Darwin’s fox(Pseudalopex fulvipes), red wolf (Canis rufus), Ethiopianwolf (C. simensis), African wild dog (Lycaon pictus), dhole(Cuon alpinus), bush dog (Speothos venaticus), Blanford’sfox (Vulpes cana) and dingo (Canis lupus dingo). In contrast,other species such as side-striped jackals (Canis adustus)and fennec foxes (Vulpes zerda) are widespread but rare(or certainly not common) throughout their range. Ourknowledge of many of the remaining species (e.g., short-eared fox, Atelocynus microtis, and pale fox, Vulpes pallida),is too poor to determine how serious are the threats theyface – although the snippets of information available offerlittle cause for optimism.

    On the other hand, many wild canids are too commonfor their own good (e.g., red fox, Vulpes vulpes, culpeo,Pseudalopex culpaeus, golden jackal, Canis aureus andcoyote, C. latrans), and thus are involved in oftencontroversial wildlife management issues (such as rabiestransmission, predation on livestock, sport hunting, furtrade).

    The canids are a fascinating family biologically(studies of them have been at the forefront of half acentury of research that has revolutionised understandingof evolutionary biology and behavioural ecology) and

    they also pose particular challenges to conservation – thetopic of this Action Plan. These two topics – the biologyand conservation of wild canids – are the subject matter ofour monograph on this family (Macdonald and Sillero-Zubiri 2004a), and many of the points that we touch onbriefly in this Introduction are fully elaborated therein.

    1.1 Canid diversity

    The number of species contained within the family Canidaeis a point of some contention, and Clutton-Brock et al.(1976), Wozencraft (1989, 1993) and Ginsberg andMacdonald (1990) argue for between 34 and 37 distinctspecies, to which the recognition of Canis lycaon (seeWilson et al. (2000) would add another. In this plan, werecognise 35 full canid species, although we discuss also a36th taxon, the dingo (Canis lupus dingo), which we considera subspecies of the grey wolf (Canis lupus).

    Canids range in size from Blanford’s (Vulpes cana) andfennec foxes, which can weigh less than 1kg, to the greywolf exceeding 60kg. Their distributions may be highlyrestricted – almost the entire Darwin’s fox populationoccurs only on one island (Yankhe et al. 1996), and someunusual subspecies occur on one island each, such asisland foxes or Mednyi Arctic foxes (Alopex lagopussemenovi – Wayne et al. 1991; Goltsman et al. 1996),

    Juvenile female Arctic fox lyingon the den at dawn. HelagsMountains, Northern Sweden,August 2002.

    Lo

    ve D

    alén

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    At least 155 of the 192 countries across the globe havecanids (81%), with Sudan the country with the highestnumber of species (10 species), followed by USA (9 species)and Ethiopia (8 species). Those countries that do not hostany canid species are island states (e.g., Caribbean islands,Madagascar, Malta, most Australasian islands).

    Africa, Asia and South America support the greatestdiversity with more than 10 canid species each. Red foxesare sympatric with 14 other canids (from three geographicalregions), golden jackals with 13 (from two regions) andgrey wolves with 11 (from three regions). Within any onelocation, however, canid diversity is usually limited to oneto five species.

    There are five canid species endemic to a single country.Not surprisingly, most are also threatened (red wolf,Ethiopian wolf, Darwin’s fox, island fox and hoary fox –Pseudalopex vetulus), with the Sechuran fox (P. sechurae)a near-endemic to Peru. Of the two continents with thehighest species diversity, South America harbours ninespecies (out of 11 species present) confined entirely tosouth of Panama, while Africa has eight endemics (of 13species present). Of 12 canid species found in Asia, onlytwo are restricted to that continent.

    Although the genera Canis and Vulpes are both foundin North America, Europe, Africa, and Asia (and wereintroduced by man to Australia), of the remaining eightgenera six are restricted to one continent: Chrysocyon,Otocyon, Pseudalopex, Speothos (South America), Cuon(Asia), Lycaon (Africa); Urocyon is restricted to Northand South America, whereas Nyctereutes, formerlyrestricted to Asia, has been introduced to Europe.

    1.3 The Canid Action Plan

    The Canid Action Plan is one in a series of such plansfostered by the World Conservation Union (IUCN) andwritten by members of the Species Survival Commission(SSC)’s Specialist Groups. The Canid Action Plan is theproduct of the deliberations of the Canid Specialist Group(CSG), itself one of more than 120 groups of specialistswith a taxonomic focus on conservation under the aegis ofSSC. The CSG has classified 36 living wild canid taxa andassigned a conservation status to each based on evaluatedrisk using the IUCN Red List Categories and Criteria:version 3.1 (IUCN 2001).

    As its name implies, The Canid Action Plan aspires toidentify important actions and to plan for theirimplementation. One way of doing this has been to canvassthe views of the international community of canid specialistson the latest knowledge and status of each species, thethreats they face, the questions that must be answered andthe actions that must be taken to ameliorate these threats.

    Paradoxically, the preparation of this Canid ActionPlan has been a unique experience precisely because we

    whereas other species span several continents – about 70million km² in the case of the red fox (Lloyd 1980). Theirdiets range from omnivory (with, at times, almost exclusiveemphasis on frugivory or insectivory) to strict carnivory –and they glean these livings in habitats ranging fromdeserts to icefields, from mountain to swamp or grassland,and from rain forest to urban ‘jungle’ (Macdonald andSillero-Zubiri 2004b). To do this they may travel homeranges as small as 0.5km² (island fox – Roemer et al. 2001)or as large and non-defensible as 2,000km² in African wilddogs (Frame et al. 1979).

    Geographical variability in body size can be explainedto some degree by differences in availability of food, withsmall canids (e.g., fennec fox) usually associated with aridand poor habitats in which only a small body mass can besupported year round, whereas large canids (e.g., Ethiopianwolf and African wild dog) are often associated withhabitats in which prey is abundant. The maned wolf(Chrysocyon brachyurus), unusual in its social organisationfor a large canid, lives in South American savannas andfeeds largely on rodents and fruit (Dietz 1985). Geffen etal. (1996) suggest that low food availability probablyconstrains both the maned wolf’s group and litter size(which is low at 2.2).

    1.2 Patterns of distribution of theCanidae

    The contemporary Canidae are the most widespread familyof extant Carnivora, with at least one species present in allcontinents except Antarctica. A quick perusal of the rangesof all canid species (Macdonald and Sillero-Zubiri 2004b)indicated that over the last century the geographical rangesof seven species have increased, eight have decreased andnine have remained stable. The kaleidoscope of speciesdiversity has changed: there are places where the grey wolfand the red fox have been replaced by what amounts totheir ‘ecological average’, the coyote (once confined tomainly arid areas in western North America and nowfound in every state, province and country north of Panama– Moore and Parker 1992; Reid 1997; Bekoff and Gese2003).

    Many Canidae have distributions that span at least awhole continent. Red foxes and grey wolves have the mostextensive natural range of any land mammal (with theexception of humans and perhaps some commensalrodents). Red foxes are the only canid present on fivecontinents, recorded in a total of 83 countries. Greywolves occur naturally in North America, Europe andAsia, their range spanning 62 countries. Two species arepresent on three continents, namely the golden jackal andArctic fox (Alopex lagopus). And two other, the red foxand dingo have reached Australia and Oceania withassistance from mankind.

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    have done it before! That conundrum is explained becausein 1990 the CSG published the first Canid Action Plan,compiled by Joshua Ginsberg and David Macdonald. Aparticularly fascinating aspect of preparing the current planhas been that it afforded us the opportunity, for the firsttime, to take stock of how things had changed and what hadbeen achieved in the intervening 14 years. We will returnbelow (section 1.4) to some lessons learnt from this longerperspective, but first we summarise the main features ofthis, the second Canid Action Plan:• This publication is an entirely new compilation of

    information on canids, reviewing exhaustively the salientadvances in science relevant to canid conservationbiology, and the most up-to-date information on statusand distribution for each species.

    • The classifications, and the proposed actions and plans,collated in this publication have been produced by themost systematic methods currently in use. To classifythe 36 taxa we have used version 3.1 of the IUCN’s RedList Categories and Criteria, and to generate candidateactions we have taken advantage of electroniccommunication to canvass a vast and internationallydispersed body of experts for their suggestions.

    • The plan includes text written by 88 contributors andreviewed by more than 90 additional referees.

    • The plan includes a comprehensive list of current projectsand researchers.

    • The entire plan will be available on the internet (http://www.canids.org), offering the facility for regular updates.

    The authors of action plans have a clear remit: to providecurrent and accurate information that will help individuals,institutions, and governments to make educated decisionswith the aim of ensuring the long-term survival of thespecies in question. To succeed in this objective, the authorsof an action plan must first collect, collate, and synthesisethe information available on the status, abundance, anddistribution of the taxon under consideration. Only thencan priorities be established and a plan for action developed.

    Some other IUCN Action Plans have organised theircontents on a regional basis, and some are further dividedinto regional reports, providing a level of detail at countrylevel that is enviable. However, such an approach is notappropriate for the canids, and we have opted to organisethe canid action plan differently. The reasons for this areseveral and stem from the basic biology of carnivores.First, carnivore species occur at lower densities than theirprey; second, at any one location, the diversity of carnivorespecies is usually rather low, while third, the geographicranges of many species are rather large. These traits ofcanid distribution are not just regional, but global. Indeciding how to present our information we were impressedby the generalisation that the status of a particular canidspecies appears to be remarkably consistent throughoutits range.

    Thus, given the wide geographical ranges of manycanid species, and their relatively low species diversity, acountry-by-country analysis of status, abundance, anddistribution seemed to us unjustifiable and potentiallyrepetitive. Worse, it would be unworkable, insofar asinformation on the status of many canids remains sparse.For most countries, one simply could not write detailedreports because detailed information is unavailable.Nonetheless, policies often have a regional focus. Manypeople involved in biological conservation are shiftingtheir attention from a species-oriented to an ecosystem-oriented approach. Therefore, in what we hope is a sensibleand utilitarian compromise between the realities of canidbiology, the limitations of the data, and the necessity fora geographical framework, we have organised this secondCanid Action Plan by species status accounts listed underrelevant geographic regions. The regions reflect, in broadterms, the biogeographical distribution of Canidae.

    1.4 The longer view

    Fifteen years have passed since the team preparing thefirst Canid Action Plan was at work and the publication ofthis second Plan. Those years have witnessed a continuingand exponential growth in understanding of, andenthusiasm for, conservation biology. Theory, far moreabundant than data in 1990, has proliferated yet faster,and with dazzling sophistication in the meantime. For thecanids as a whole, empirical knowledge has, if anything,fallen yet further behind – the corsac fox, the small-eareddog and the Indian fox are still as unknown as they werein 1990 – but the situation is partly saved because theintervening years have seen the publication of somewonderful field studies (and 14 of these are highlighted ascase studies in Macdonald and Sillero-Zubiri 2004a).Despite the gaps, therefore, a great deal more is knownnow than then. Against that background, revisiting thefirst plan while working on this second one has caused usto ask what has changed. Some things have not.

    Foremost amongst the things that have not changedare the inherent natures of wild canids and of people –natures that all too often throw us into conflict. Thus, it isstill true that the fox trotting across your field of view maybe, simultaneously, a resource for the trapper, a healthhazard to the rabies official, a quarry to the huntsman, asubject to the photographer, vermin to the poultry farmer,and a joy to behold to the aesthete. What is more, it is stilltrue that their judgements are neither right nor wrongaccording to some self-appointed prophet – each couldargue a case, but would do so using incommensurablecurrencies (how are we to equate units of jobs versus unitsof cultural heritage versus units of suffering versus money,etc.?). Again and again, scientific judgement trips overethical judgement. In the Introduction to the first Canid

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    Action Plan we wrote of the challenge of untangling theseissues as the prerequisite to resolving them – it remains noless a challenge now.

    Science has been helpful. For example, the oralvaccination of foxes against rabies was avant garde in the1980s, but is de rigour now. On the other hand, alternative,and often non-lethal, approaches to resolving conflictswith canids that we vaunted expectantly in 1990 – such asaversive conditioning or sophisticated repellents – havestill not really materialised (although the scope is great; seeSillero-Zubiri et al. (2004)). On the other hand, appreciationof the human dimension to conservation biology hasadvanced hugely, and the processes and outcomesassociated with the reintroduction of wolves to Yellowstoneillustrate the new deal. In this vein, exciting, collaborative,often community-based projects are developing, and thetrial introduction of African wild dogs to a consortia ofadjoining South African game ranches is a case in point:the question is whether some way can be found to ensurethat the profit from ecotourists attracted by the wild dogswill outweigh the losses caused by their depredations onvaluable antelope – the research is underway and theanswer may be some years in coming, but it is exciting thatthe question is at least being both asked and tackled.

    In 1990 we wrote that the very opportunism and mobilitythat is the hallmark of success for many members of the dogfamily is also responsible for throwing them into conflictwith people. This conflict involves, first, competitionbetween man and canids through depredations on gameand domestic stock; and, second, canids as victims of, andoften vectors for, several zoonoses, of which rabies isundeniably the most notable. In addition to rabies, thereare several other pathogens of direct (e.g., leishmaniasis)and indirect (e.g., sarcoptic mange) concern to man, thelatter threatening the fur trade, the former threatening life.The traditional response to perceived problems of predationand disease has been to attempt to reduce canid numbersby killing them. This time-honoured approach had, andstill has, two notable drawbacks: it tends to throw differentfactions into conflict (e.g., fur traders object to theircommodity being blasted by irate shot-gun-totingstockmen, and those concerned with animal welfare objectto both groups killing canids); and it may not work (eitherbecause the problem was merely perceived but not real, orbecause the solution was inappropriate). The issues raisedby predation and disease are biologically and economicallycomplicated. Data on both topics are more revealingnowadays, and Part 3 of this Action Plan considers thesecontinuing realities within a much more secure frameworkof ideas than was available in the first edition.

    Our intention remains to identify problems commonto many canids, real and imaginary, to seek some indicationof their magnitudes, and to evaluate the effectiveness ofexisting solutions and the practicability of novel ones, allfrom the point of view of species conservation. Perhaps

    the most clear-cut change in attitude is that disease hasbecome accepted as a two-faced demon in canidconservation: not only is infectious disease a scourge ofpopulous canids, and thus a threat to mankind and hisdomestic stock, but it is also a threat to rare canidsthemselves – sometimes because numerous canids threatenthem (uncommon Blanford’s foxes might succumb torabies transmitted by abundant red foxes), but morecommonly because rare wild canids (Ethiopian wolves,African wild dogs, island foxes) are decimated by diseasesof domestic dogs. Happily, this realisation has fosteredproductive links between biologists, veterinarians, publichealth officials and community development experts. Thisis the sort of interdisciplinarity that we hailed as necessaryin 1990, and which is now widely becoming familiar – butthere is still a long way to go.

    1.5 Structure of the Action Plan

    The arrangement of species sections within chapters followsa regional approach. Each of the world’s canid species hasbeen assigned to one of seven regional chapters (Chapters3–9). The structure of each geographically organisedchapter is explained in the Introduction of Part 2 “SpeciesStatus Accounts”.

    Our approach was to seek more than one author foreach entry and then to support them with a team ofreviewers. The reviewers were selected, where possible, tospan the geographic region covered by the species inquestion. The final entry, therefore, combines the expertiseof the authors, reviewers and editors in what we believeare the most comprehensive accounts available. A firststep in this process, however, was to agree which speciesare to be recognised, and by which names are we to referto them. Therefore, in Chapter 2 there is a discussion ofcanid taxonomy, and the classification used in thispublication is explained. Although the Action Plan doesnot attempt to analyse conservation priorities at the levelof subspecies (with the exception of the dingo, hereinconsidered a subspecies of the grey wolf), each speciesaccount includes a list of recognised subspecies for thosewho wish to pursue questions of taxonomy andbiogeography.

    Foxes, wolves, and jackals have much in common, bothin their biology, their relationships with man and theirconservation challenges. Hence, following the geographicalchapters, in Part 3 we present nine chapters on topicsalmost universally relevant to the conservation of theCanidae as a whole. The topics are sociobiology, genetics,diseases, management of canids near people, trade, researchtechniques, captive breeding, reintroduction, conservationpolicies, and conservation education.

    Finally, Part 4 represents the true ‘action’ componentof the plan, an ‘Action Plan for Canid Conservation in the

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    21st century’, introduced by a short chapter on the increasingneed in conservation biology for setting priorities andmeasuring success (Chapter 19). In chapter 20 we list theprojects and actions that we believe are priorities for canidconservation over the next 10 years. These are presented ina simple summary form. Given that canids are presentthroughout the world, and that many of the range countriesdo not have appropriate scientific and conservationinfrastructure, we have attempted to identify the mostneeded projects and actions to improve the conservationstatus of canid species, rather than present a “wish” list ofprojects per country as other IUCN action plans havedone. We aim to achieve a realistic list of projects andactions that would have a good chance of being implementedwithin a decade. Thus the projects/actions focus on thethreatened species, and are organised into two sections; a)general projects/actions affecting all species, and b) projects/actions listed under relevant species. The process ofassembling these proposals was revealing, as was the natureof the proposals themselves – we feel certain generalisationsemerge from them, and we have discussed these at length inMacdonald and Sillero-Zubiri (2004c).

    During the three years or so that it has taken toproduce this Action Plan we received many reports fromCSG members and other canid enthusiasts. We receivedstandardised questionnaire forms providing informationon the distribution and status of wild canids at countrylevel. Such information was made available to the authorsof each species’ account. The second Canid Action Plancould not have been completed without the help of thesecorrespondents, and we have tried to represent their viewsaccurately. Many of our correspondents have emphasisedthe potential usefulness of a register of conservation andresearch projects dealing with canids and those peopleinvolved with canid research. This has prompted the CSGto develop a web-based Canid Project Database (http://www.canids.org/database/ – see Appendix 5). Thisdirectory is by no means complete, and we encouragereaders to submit information on those projects that arenot yet covered by the database.

    1.6 Limitations of this Action Plan

    The structure we have imposed upon this second editionof the Canid Action Plan has suited our purpose, but it hasdrawbacks. For example, by making our focus regionaland global, we have largely ignored aspects of conservationat the level of subspecies or of local populations. Indefence of this, we would argue that, for the most part,patterns of extinction are regional: a succession ofsubpopulations disappears, survivors become fragmented,and local extirpations start the slippery slide to extinction.In addition, for many species such as the short-eared dog(Atelocynos microtis) or the pale fox (Vulpes pallida), data

    are so inadequate that the available materials swiftlydetermined the scope of our accounts.

    The nomination of priorities also lures us into animponderable mirage dividing biology and ethics. It maybe tempting to seek criteria on which to decide whether itis more important to save the Ethiopian wolf than the redwolf. But both are irreplaceable. We might be able to guessthe order in which threatened canid species will go extinct,but we have tried not to place relative value on the loss ofone versus another. Nonetheless, as funds and time arelimited, we have tried to focus our action planning onthose species most threatened with imminent extinction,and secondly those for which more data are needed inorder to make judgement.

    1.7 The Canid Specialist Group

    The Canid Specialist Group (CSG) is the world’s chiefbody of scientific and practical expertise on the statusand conservation of all species of the Canidae (wolf,jackal and fox family), advising the Species SurvivalCommission (SSC) of the World Conservation Union(IUCN). The CSG is based at the Wildlife ConservationResearch Unit (WildCRU), Oxford University, UK (http://www.wildcru.org).

    The CSG is composed of more than 100 experts,representing over 30 countries (and with experience inmany more). These people serve as honorary advisors tothe CSG in their personal capacities, but bring with themthe experience and the knowledge gained in theirprofessional careers. CSG membership is rapidlyexpanding and is open to anybody actively involved incanid conservation and research. There is a separate WolfSpecialist Group concerned specifically with grey and redwolves.

    The Mission of the CSG is to promote the long-termconservation of all Canid species throughout their ranges

    Objectives:• Compile, synthesise and disseminate information on

    the conservation of all canids across their range, withparticular emphasis on species that are threatened orrare.

    • Provide technical information and advice on all mattersconcerning wild canids, including their status in thewild; their biology and natural history; the threats theyface and their conservation requirements to thefollowing:a) range state government agencies;b) national and international NGOs, including

    potential funding bodies;c) inter-governmental organisations (e.g., IUCN,

    CITES);d) field projects concerned with canid conservation.

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    • Promote and catalyse conservation activities on behalfof wild canids, prioritising and coordinating efforts ofresearchers and conservationists worldwide.

    • Fundraise for canid research and conservation andundertake research directly when necessary orappropriate.

    • Improve management of the common and sometimestroublesome species.

    • Build capacity through the exchange of ideas,information, and technical expertise among themembers of the Group.

    CSG functions and activities• Liaison between field biologists, wildlife managers,

    governments, NGOs and sponsors on any topicconcerning the conservation of canid species.

    • Compilation, synthesis and dissemination of canidrelated information.

    • Development of the Canid Conservation Database, anupdated list of references and information on currentresearch and conservation projects.

    • Hold regular meetings of CSG members alongsideinternational conferences.

    • Serve as the IUCN Red List Authority for the Canidae,responsible to evaluate the category of threat of allcanids.

    • Regionalised approach – A global network withRegional Section Chairs for: Sub-Saharan Africa;North Africa and Middle East; North and CentralAsia; South Asia and Australasia; Europe; North andCentral America; South America.

    • Species Working Groups – Bringing together theexperts of a particular species: African wild dog;Arctic fox; Ethiopian wolf; dhole; island fox, kit foxand swift fox.

    • Topical Working Groups – Addressing specificproblems across species: Disease and Epidemiology;Ecology and Research; Genetics; Harvesting and PestControl; International Policy; Re-introduction andTranslocation.

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    The family Canidae belongs to the order Carnivora, a largegroup of mostly predatory mammals characterised bytheir common possession of a pair of carnassial teeth(upper fourth premolar and lower first molar) that aremodified to maximise efficiency for shearing skins, tendons,and muscles in their preys. Canids are characterised by aninflated entotympanic bulla (bony chamber enclosing themiddle ear region) that is divided by a partial septum alongthe entotympanic and ectotympanic suture. Other featurescharacteristic of canids are the loss of a stapedial artery andthe medial position of the internal carotid artery that issituated between the entotympanic and petrosal for mostof its course and contained within the rostral entotympanicanteriorly (Wang and Tedford 1994). These basicranialcharacteristics have remained more or less stable throughoutthe history of canids, allowing easy identification in thefossil record when these structures are preserved.

    2.1 Phylogeny from morphologicaland palaeontological perspective

    There are three major groups (subfamilies) in the familyCanidae: Hesperocyoninae, Borophaginae and Caninae(Tedford 1978) (Figures 2.1, 2.2). Of these, two arerepresented by fossil forms only. The Hesperocyoninae isthe most ancient group of all canids, and its basal member,Hesperocyon, gave rise to the two more advancedsubfamilies, Borophaginae and Caninae (Wang 1994). Amajor evolutionary transformation involves themodification of the talonid heels on the lower carnassialtooth (first lower molar), which changes from that of atrenchant, blade-like condition in the Hesperocyoninae tothat of a basined condition enclosed by two cusps in theBorophaginae and Caninae (Figure 2.3). Mainly due totheir common possession of this basined talonid, theBorophaginae and Caninae are hypothesised to share acommon ancestry. Along with a more quadrate upper firstmolar with a large hypocone on the inner corner of thetooth, the basined talonid establishes an ancestral statefrom which all subsequent forms were derived. Such adental pattern has proved to be very versatile and can

    readily be adapted toward either a highly carnivorous ora less carnivorous type of dentition, both of which wererepeatedly employed by both Borophaginae and Caninae.

    The extinct Borophaginae was the first major group ofcanids to demonstrate the viability of a basined talonidand achieved the greatest morphological breadth andtaxonomical diversity within the North Americancontinent (Wang et al. 1999). Toward the less predaceousend of the morphological spectrum, it sports highlyomnivorous forms that parallel similar adaptations byliving Procyonidae (the raccoon family). Toward the morepredaceous end, on the other hand, the Borophaginae iswell known for its tendencies to develop strong bone-crushing dentitions that parallel the habits of livingHyaenidae (the hyaena family).

    The subfamily Caninae started with Leptocyon, anancestral species the size of a small fox. Besides sharing abicuspid talonid of M1 and a quadrate M1 with theborophagines, Leptocyon is also characterised by a slenderrostrum and elongated lower jaw, and correspondinglynarrow and slim premolars, features that are inherited inall subsequent canines. It first appeared in the earlyOligocene (Orellan, 34–32 million years before present[BP]) and persisted through the late Miocene(Clarendonian, 12–9 million years BP). Throughout itslong existence (no other canid genus had as long a duration),facing intense competition from the larger and diversehesperocyonines and borophagines, Leptocyon generallyremained small in size and low in diversity, never havingmore than two or three species at a time.

    By the latest Miocene (Hemphillian, 9–5 million yearsBP), fox-sized niches are widely available in NorthAmerica, left open by extinctions of all small borophagines.The true fox clade, Tribe Vulpini, emerges at this time andundergoes a modest diversification to initiate primitivespecies of both Vulpes and Urocyon (and their extinctrelatives). The North American Pliocene record of Vulpesis quite poor. Fragmentary materials from early Blancanindicate the presence of a swift fox (Vulpes velox) in theGreat Plains. Vulpes species were widespread and diversein Eurasia during the Pliocene (see Qiu and Tedford 1990),resulting from an immigration event independent from

    Chapter 2

    Phylogeny, Classification, and EvolutionaryEcology of the Canidae1

    Wang Xiaoming, R.H. Tedford, B. Van Valkenburgh, and R.K. Wayne

    1 This paper is adapted from Wang, X., Tedford, R.H., Van Valkenburgh, B. and Wayne, R.K. 2004. Evolutionary history, molecular systematics, and evolutionary ecology of Canidae.Pp. 38–54 in D.W. Macdonald and C. Sillero-Zubiri (eds). Biology and conservation of wild canids, Oxford University Press, Oxford, UK.

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    Figure 2.1. Simplified phylogenetic relationships of canids at the generic level.Species ranges are indicated by individual bars enclosed within grey rectangles, detailed relationships among species in a genus is notshown. Relationships for the Hesperocyoninae is modified from Wang (1994: fig. 65), that for the Borophaginae from Wang et al. (1999:fig. 141), and that for the Caninae from unpublished data by Tedford, Wang, and Taylor.

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    Figure 2.2. Dental evolution of representative canids as shown in upper cheek teeth (P4-M2).Generally the most derived species in each genus is chosen to enhance a sense of dental diversity. Species in the Hesperocyoninaeare: Hesperocyon gregarius; Paraenhydrocyon josephi; Cynodesmus martini; Enhydrocyon crassidens; and Osbornodon fricki. Species inthe Borophaginae are: Cynarctoides acridens; Phlaocyon marslandensis; Desmocyon thomsoni; Cynarctus crucidens; Euoplocyonbrachygnathus; Aelurodon stirtoni; Paratomarctus temerarius; Carpocyon webbi; Epicyon haydeni; and Borophagus diversidens. Speciesin the Caninae are: Leptocyon gregorii; Vulpes stenognathus; Urocyon minicephalus; Cerdocyon thous; Eucyon davisi; Canis dirus; andCuon alpinus. All teeth are scaled to be proportional to their sizes.

    Figure 2.3. Hypercarnivorous (B, Aelurodon andD, Euoplocyon) and hypocarnivorous (A, Phlaocyonand C, Cynarctus) dentitions.In hypercarnivorous forms, the upper cheek teeth (B) tend toemphasise the shearing part of the dentition with an elongated andnarrow P4, an enlarged parastyle on a transversely elongated M1,and a reduced M2. On the lower teeth (D), hypercarnivory isexemplified by a trenchant talonid due to the increased size andheight of the hypoconid at the expense of the entoconid (reducedto a narrow and low ridge), accompanied by the enlargement of theprotoconid at the expense of the metaconid (completely lost inEuoplocyon) and the elongation of the trigonid at the expense of thetalonid. In hypocarnivorous forms, on the other hand, the upperteeth (A) emphasise the grinding part of the dentition with ashortened and broadened P4 (sometimes with a hypocone alongthe lingual border), a reduced parastyle on a quadrate M1 that hasadditional cusps (e.g., a conical hypocone along the internalcingulum) and cuspules, and an enlarged M2. The lower teeth (C) inhypocarnivorous forms possess a basined (bicuspid) talonid on m1enclosed on either side by the hypoconid and entoconid that areapproximately equal in size. Other signs of hypocarnivory on thelower teeth include widened lower molars, enlarged metaconids,and additional cuspules such as a protostylid.

    MaMa

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    that of the Canis clade. Red fox (Vulpes vulpes) and Arcticfox (Alopex lagopus) appeared in North America only inthe late Pleistocene, evidently as a result of immigrationback to the New World.

    Preferring more wooded areas, the gray fox (Urocyon)has remained in southern North America and CentralAmerica. Records of the gray fox clade have a more or lesscontinuous presence in North America throughout itsexistence, with intermediate forms leading to the livingspecies U. cinereoargenteus. Morphologically, the livingAfrican bat-eared fox (Otocyon megalotis) is closest to theUrocyon clade, although molecular evidence suggests thatthe bat-eared fox lies at the base of the fox clade or evenlower (Geffen et al. 1992; Wayne et al. 1997). If themorphological evidence is correct, then the bat-eared foxmust represent a Pliocene immigration event to the OldWorld independent of other foxes.

    Advanced members of the Caninae, Tribe Canini, firstoccur in the medial Miocene (Clarendonian) in the form ofa transitional taxon Eucyon. As a jackal-sized canid,Eucyon is mostly distinguished from the Vulpini in anexpanded paroccipital process and enlarged mastoidprocess, and in the consistent presence of a frontal sinus.The latter character initiates a series of transformations inthe Tribe Canini culminating in the elaborate developmentof the sinuses and a domed skull in the grey wolf (Canislupus). By the late Miocene, species of Eucyon haveappeared in Europe (Rook 1992) and by the early Pliocenein Asia (Tedford and Qiu 1996). The North Americanrecords all predate the European ones, suggesting awestward dispersal of this form.

    Arising from about the same phylogenetic level asEucyon is the South American clade. Morphological andmolecular evidence generally agrees that living SouthAmerican canids, the most diverse group of canids on asingle continent, belong to a natural group of their own.The South American canids are united by morphologicalcharacters such as a long palatine, a large angular processof the jaw with a widened scar for attachment of theinferior branch of the medial pterygoid muscle, and arelatively long base of the coronoid process (Tedford et al.1995). By late Hemphillian and early Blancan, certainfragmentary materials from southern United States andMexico indicate that the earliest taxa assignable toCerdocyon (Torres and Ferrusquía-Villafranca 1981) andChrysocyon occur in North America. The presence ofthese derived taxa in the North American late Miocenepredicts that ancestral stocks of many of the SouthAmerican canids may have been present in southern NorthAmerica or Central America. They appear in the SouthAmerican fossil record shortly after the formation of theIsthmus of Panama in the Pliocene, around three millionyears BP (Berta 1987). The earliest records are Pseudalopexand its close relative Protocyon, an extinct largehypercarnivore, from the Pliocene (Uquian, around 2.5–

    1.5 million years BP) of Argentina. By the late Pleistocene(Lujanian, 300,000–10,000 years BP), most living speciesor their close relatives have emerged, along with theextinct North American dire wolf (Canis dirus). By the endof the Pleistocene, all large, hypercarnivorous canids ofSouth America (Protocyon, Theriodictis) as well as Canisdirus had become extinct.

    The Canis-Lycaon clade within the Tribe Canini, themost derived group in terms of large size andhypercarnivory, arose near the Miocene-Plioceneboundary between 6 and 5 million years BP in NorthAmerica. A series of jackal-sized ancestral species ofCanis thrived in the early Pliocene (early Blancan), such asC. ferox, C. lepophagus, and other undescribed species. Atabout the same time, first records of canids begin toappear in the European late Neogene: Canis cipio in thelate Miocene of Spain (Crusafont-Pairó 1950), Eucyonmonticinensis in the late Miocene of Italy (Rook 1992), theearliest raccoon-dog (Nyctereutes donnezani), and thejackal-sized Canis adoxus in the early Pliocene of France(Martin 1973; Ginsburg 1999). The enigmatic C. cipio,only represented by parts of the upper and lower dentitionat a single locality, may represent a form at the Eucyonlevel of differentiation rather than truly a species of Canis.

    The next phase of Canis evolution is difficult to track.The newly arrived Canis in Eurasia underwent an extensiveradiation and range expansion in the late Pliocene andPleistocene, resulting in multiple, closely related species inEurope, Africa and Asia. To compound this problem, thehighly cursorial wolf-like Canis species apparently belongto a circum-arctic fauna that undergoes expansions andcontractions with the fluctuating climates.Hypercarnivorous adaptations are common in the crown-group of species especially in the Eurasian middle latitudesand Africa. For the first time in canid history, phylogeneticstudies cannot be satisfactorily performed on forms fromany single continent because of their Holarctic distributionand faunal intermingling between the New and Old Worlds.Nevertheless, some clades are localised in different partsof the Holarctic. The vulpines’ major centre of radiationwas in the Old World. For the canines, North Americaremained a centre through the Pliocene producing thecoyote (Canis latrans) as an endemic form. The wolves,dhole (Cuon alpinus), African wild dog (Lycaon pictus)and fossil relatives are the products of the Eurasian andAfrican continents. During the Pleistocene, elements ofthe larger canid fauna invaded mid-latitude North America– the last invasion of which was the appearance of the greywolf south of the glacial ice sheets in the late Pleistocene(about 100,000 years BP).

    A comprehensive systematic revision of NorthAmerican fossil canines by Tedford et al. (in prep.) is nearcompletion, which forms the basis of much of the abovesummary. As part of the above revision, the phylogeneticframework as derived from living genera was published by

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    Tedford et al. (1995). Nowak (1979) monographed theQuaternary Canis of North America; Berta (1981, 1987,1988) did the most recent phylogenetic analysis of theSouth American canids; Rook (1992, 1994) and Rook andTorre (1996a, 1996b) partially summarised the Eurasiancanids. The African canid records are relatively poorlyunderstood and recent discoveries promise to significantlyadvance our knowledge in that continent (L. Werdelinpers. comm.).

    2.2 Molecular phylogeny

    The ancient divergence of dogs from other carnivores isreaffirmed by molecular data. DNA-DNA hybridisationof single copy DNA clearly shows them as the firstdivergence in the suborder Caniformia that includespinnipeds, bears, weasel and raccoon-like carnivores(Figure 2.4). This basal placement is further supported bymitochondrial DNA (mtDNA) sequence studies (Vranaet al. 1994; Slattery and O’Brien 1995; Flynn and Nedbal1998). Based on molecular clock calculations, thedivergence time was estimated as 50 million years BP(Wayne et al. 1989). This value is consistent with the firstappearance of the family in the Eocene, although it issomewhat more ancient than the date of 40 million yearssuggested by the fossil record (see above). Consideringthat first appearance dates generally postdate actualdivergence dates because of the incompleteness of therecord (e.g., Marshall 1977), the agreement between fossiland molecular dates is surprisingly good.

    Evolutionary relationships within the family Canidaehave been reconstructed using comparative karyology,allozyme electrophoresis, mtDNA protein coding sequencedata, and, most recently, supertree method (Wayne andO’Brien 1987; Wayne et al. 1987a, 1987b, 1997; Bininda-Emonds et al. 1999). Further, relationships at the genuslevel have been studied with mtDNA control regionsequencing (a non-coding, hypervariable segment of about1200 years BP in the mitochondrial genome) andmicrosatellite loci (hypervariable single copy nuclear repeatloci) (Geffen et al. 1992; Bruford and Wayne 1993; Girmanet al. 1993; Gottelli et al. 1994; Vilà et al. 1997, 1999). Theprotein-coding gene phylogeny, which is largely consistentwith trees based on other genetic approaches, shows thatthe wolf genus Canis is a monophyletic group that alsoincludes the dhole or Asian wild dog. The grey wolf,coyote and Ethiopian Wolf or Simien Jackal (Canissimensis) form a monophyletic group, with the goldenjackal (C. aureus) as the most likely sister taxon (Figure2.5). The black-backed (C. mesomelas) and side-stripedjackals (C. adustus) are sister taxa, but they do not form amonophyletic group with the golden jackal and Ethiopianwolf. Basal to Canis and Cuon are the African wild dogand a clade consisting of two South American canids, the

    Figure 2.4. Relationship of carnivores based onDNA hybridisation data (Wayne et al. 1989).Family and suborder groupings are indicated. Time scale inmillions of year before present (MYBP) is based on comparisonsof DNA sequence divergence to first appearance times in thefossil record.

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    bush dog (Speothos venaticus) and the maned wolf(Chrysocyon brachyurus). Consequently, although theAfrican wild dog preys on large game as does the grey wolfand dhole, it is not closely related to either species but issister to the clade containing these species. This phylogenyimplies that the trenchant-heeled carnassial now foundonly in Speothos, Cuon and Lycaon, evolved at least twiceor was primitive and lost in other wolf-like canids and themaned wolf.

    The South American canids do not form a monophyleticgroup. Speothos and Chrysocyon are sister taxa that groupwith the wolf-like canids rather than the South Americanfoxes. The large sequence divergence between the bush

    dog and maned wolf and between these taxa and the SouthAmerican foxes suggests that they diverged from eachother 7–6 million years BP, well before the Panamanianland bridge formed about 3–2 million years BP. Thus,three canid invasions of South America are required toexplain the phylogenetic distribution of the extant species.These invasions are today survived by the bush dog,maned wolf, and the South American foxes. Further,within the South American foxes, divergence valuesbetween crab-eating fox (Cerdocyon thous), the short-eared dog (Atelocynus microtis) and other South Americanfoxes, suggest they may have diverged before the openingof the Panamanian land bridge as well (Wayne et al. 1997).

    Figure 2.5. Consensustree of 26 canidspecies based onanalysis of 2,001 bp ofDNA sequence frommitochondrial proteincoding genes (Wayneet al. 1997).See Geffen et al. (1992) for amore detailed analysis of theRed-fox like canids. Time scalein millions of year beforepresent (MYBP) is based oncomparisons of DNA sequencedivergence to first appearancetimes in the fossil record.

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    The fossil record supports the hypothesis that the crab-eating fox had its origin outside of South America as thegenus has been described from late Miocene deposits ofNorth America (6–3 million years BP) (Berta 1984, 1987,see above). Consequently, only the foxes of the genusPseudalopex, Lycalopex and perhaps Atelocynus, appearto have a South American origin. Further, the genericdistinction given to Pseudalopex and Lycalopex does notreflect much genetic differentiation, and in the absence ofappreciable morphologic differences, the genetic datasuggest these species should be assigned to a single genus.

    A fourth grouping in the tree consists of other fox-liketaxa, including Alopex and Vulpes (here considered toinclude the fennec fox, Vulpes zerda, sometimes includedin the genus Fennecus) (Figure 2.5) (Geffen et al. 1992;Mercure et al. 1993; Wayne et al. 1997). The Arctic fox isa close sister to the kit fox (Vulpes macrotis) and bothshare the same unique karyotype (Wayne et al. 1987a).Finally, Otocyon, Nyctereutes, and Urocyon appear basal

    to other canids in all molecular and karyological trees(Wayne et al. 1987a). The first two taxa are monospecificwhereas the third includes the island fox (Urocyon littoralis)and the gray fox (U. cinereoargenteus). The three generadiverged early in the history of the family, approximately12–8 million years BP as suggested by molecular clockextrapolations.

    In sum, the living Canidae is divided into five distinctgroupings. These include the wolf-like canids, whichconsists of the coyote, grey wolf, Ethiopian wolf, jackals,dhole and African wild dog. This clade is associated witha group containing bush dog and maned wolf in some treesand, further, this larger grouping is associated with theSouth American foxes (Wayne et al. 1997). The red foxgroup is a fourth independent clade containing Alopexand Vulpes (including the fennec fox). Finally, threelineages have long distinct evolutionary histories and aresurvived today by the raccoon dog, bat-eared fox andisland and gray fox. Assuming an approximate molecular

    Yearling male island fox priorto dispersal from natal area.Fraser Point, Santa CruzIsland, California, USA, 1993.

    Gar

    y R

    oem

    er

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    clock, the origin of the modern Canidae begins about 12–10 million years BP and is followed by the divergence ofwolf and fox-like canids about 6 million years BP. TheSouth American canids are not a monophyletic group andlikely owe their origin to three separate invasions. Thisgroup included the maned wolf, bush dog, crab-eating foxand the other South American canids that diverged fromeach other about 6–3 million years BP.

    2.3 Evolutionary ecology

    2.3.1 Iterative evolution ofhypercarnivory

    One of the most remarkable features of canid history istheir repeated tendency to evolve both hypocarnivorousand hypercarnivorous forms. As noted above,hypercarnivorous species evolved within each subfamily,and hypocarnivorous species evolved within two of thethree (all but the Hesperocyoninae). Hypocarnivory wasmost fully expressed in the Borophaginae, where at least15 species showed a tendency towards a dentition similarto that of living raccoons (Wang et al. 1999). Among theCaninae, the tendency has not been quite as strong, withonly a single lineage, Nyctereutes, developing a markedlyhypocarnivorous dentition. However, all three subfamiliesinclude multiple species of apparent hypercarnivores withenhanced cutting blades on their carnassials, reducedgrinding molars, and enlarged canines and lateral incisors.When and why did hypercarnivory evolve within eachsubfamily?

    In two of the three subfamilies, Hesperocyoninae andCaninae, the evolution of hypercarnivory appears to haveoccurred at least partly in response to a reduced diversityof other hypercarnivorous taxa. The Hesperocyoninaeevolved hypercarnivory early in their history (Figures 2.1,

    2.2, 2.6) and the most advanced forms appear in the earlyMiocene (about 24–20 million years BP) at a time whenthe two previously dominant carnivorous families hadvanished. These two families were the Nimravidae, anextinct group of saber-tooth cat-like forms, and theHyaenodontidae, a group of somewhat dog-like predatorsincluded in the extinct order Creodonta. The nimravidsand hyaenodontids dominated the North American guildof large, predatory mammals in the late Eocene to mid-Oligocene (37–29 million years BP), but faded rapidly inthe late Oligocene, and were extinct in North America byabout 25 million years BP (Van Valkenburgh 1991, 1994).During most of their reign, hesperocyonines existed at lowdiversity and small (fox-size) body size, but as thehyaenodontids and nimravids declined in the lateOligocene, the early canids seem to have radiated toreplace them. Most of these hypercarnivorous canidswere jackal-size (less than 10kg), with only the last survivingspecies, Osbornodon fricki, reaching the size of a smallwolf (Wang 1994). In the early Miocene, largehypercarnivores emigrated from the Old World in theform of hemicyonine bears (Ursidae) and temnocyoninebear-dogs (Amphicyonidae). The subsequent decline toextinction of the hesperocyonines might have been a resultof competition with these new predators (Van Valkenburgh1991, 2001).

    Hypercarnivory appears late in the history of theCaninae and represents at least several independentradiations in South America, North America, and the OldWorld (Figures 2.1, 2.6). As was true of the hesperocyonineexample, the South American radiation of largehypercarnivorous canids occurred at a time (2.5–0.01million years BP) when cat-like predators were rare orabsent. It followed the elevation of the Panamanian landbridge around 3–2 million years BP that allowedimmigration between the previously separated continents.The canids that first entered South America found a

    Figure 2.6. Iterative evolution of large hypercarnivores.Number (N) of hypocarnivorous (white), mesocarnivorous (grey), and large (>20kg) hypercarnivorous (black) species over time in each ofthe three subfamilies. The few hesperocyonine species with trenchant-heeled carnassials estimated to have been less than 20kg in masswere assigned to the mesocarnivorous category because they are assumed not have taken prey as large or larger than themselves. Forthe Hesperocyoninae and Borophaginae, their stratigraphic ranges were broken into thirds; for the Caninae, four time divisions wereused because of the large number of species appearing in the past five million years. Species were assigned to dietary categories andbody mass was estimated on the basis of dental morphology as described in Van Valkenburgh (1991) and Wang et al. (1999).

    Hesperocyoninae Borophaginae Caniinae

    Millions of years before presentMillions of years before present Millions of years before present

    N N N

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    depauperate predator community, consisting of one bear-like procyonid carnivoran, three species of carnivorousdidelphid marsupials, one of which was the size of acoyote, and a gigantic, predaceous ground bird (Marshall1977). With the possible exception of the rare ground bird,none of these species was a specialised hypercarnivore.Between 2.5 million and 10,000 years BP, 16 new speciesof canids appeared in South America, at least seven ofwhich had trenchant-heeled carnassials and clearly wereadapted for hypercarnivory (Berta 1988; Van Valkenburgh1991). They represent three different endemic genera:Theriodictis, Protocyon and Speothos. In addition, therewere three large wolf-like species of Canis in SouthAmerica, C. gezi, C. nehringi, and C. dirus, all of whichwere probably hypercarnivo