foxes, wolves, jackals, and dogs - international union for ... · foxes, wolves, jackals, and dogs...

of 123/123
Foxes, Wolves, Jackals, and Dogs An Action Plan for the Conservation of Canids J.R. Ginsberg and D.W. Macdonald IUCN/SSC Canid Specialist Group IUCN/SSC Wolf Specialist Group (L.D. Mech, Chair)

Post on 27-Apr-2018

263 views

Category:

Documents

3 download

Embed Size (px)

TRANSCRIPT

  • Foxes, Wolves, Jackals,and Dogs

    An Action Plan for the Conservation of Canids

    J.R. Ginsberg and D.W. Macdonald

    IUCN/SSC Canid Specialist GroupIUCN/SSC Wolf Specialist Group (L.D. Mech, Chair)

  • This publication has been produced by IUCNThe World Conservation Union, in collaboration with the International Fur Trade Federation (IFTF), the WorldWide Fund for NatureInternational (WWF), the People's Trust for Endangered Species (PTES), the American Association of Zoological Parks and Aquariums(AAZPA), and the Oxford Wildlife Conservation Research Unit (WCRU).

    1990 International Union for Conservation of Nature and Natural Resources

    Reproduction of this publication for educational and other non-commercial purposes is authorized without permission from the copyright holder, provided thesource is cited and the copyright holder receives a copy of the reproduced material.

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

    ISBN 2-88032-996-5

    Published by: IUCN, Gland, Switzerland

    Printed by: Kelvyn Press, Inc., Broadview, Illinois, United States of America

    Cover photo: African wild dog (Lycaon pictus). (Photo by J.R. Ginsberg)

  • Foxes, Wolves, Jackals,and Dogs

    An Action Plan for the Conservation of Canids

    J.R. Ginsberg and D.W. Macdonald

    IUCN/SSC Canid Specialist GroupIUCN/SSC Wolf Specialist Group (L.D. Mech, Chair)

  • Contents

    Page

    Forewords and Acknowledgements iii

    Section 1. Introduction 1

    1. Objectives, Structure, and Limitationsof the Canid Action Plan 1

    2. Sorting Out the Canidae 4

    Section 2. Species and Geographic Analyses 10

    3. Introduction to Species Summaries andGeographical Chapters 10

    4. Sub-Saharan Africa 11

    5. South America 23

    6. Holarctic 33

    7. North Africa and the Middle East 48

    8. South and Southeast Asia and Australia 52

    Page

    Section 3. General Considerations 58

    9. International Trade in CITES

    Listed Canids 58

    10. Captive Breeding and Reintroduction 68

    11. Predation and Predator Control 77

    12. Canids and Disease 86

    13. Conclusions and Action Planning 89

    Appendices 102

    1. List of the Members of the IUCN/SSCCanid Specialist Group 102

    2. Other Correspondents Who Contributedto the Compilation of the Canid Action Plan 103

    3. List of Members of the IUCN/SSCWolf Specialist Group 109

    References 110

    Forewords and Acknowledgements

    Perhaps it is insight into their individualism won through ourassociation with domestic dogs, perhaps it is the ability of someof them to withstand the worst that man can throw at them,perhaps it is that we see some reflection of ourselves in theiropportunism, or perhaps it is simply the transfixing elegance oftheir jaunty grace. Whatever the pot-pourrie of reasons thatblend into their appeal, the 34 species that comprise the familyCanidae are special. They are special because they have, asperceived friend or foe, preoccupied the imaginations of man-kind for millennia. They are special because the breadth of theiradaptations makes them enthralling to science. They are specialbecause the contradictory facets of their relations with peopleperplex the conservationist. The possibility that we are heed-lessly, perhaps needlessly, mismanaging many of them is sad-dening; the probability that our negligence will force severalmore to extinction should fill us with bottomless dismay. Itdemands action, and that is why we have written the CanidAction Plan.

    Species action plans are produced by the Specialist Groupsof the Species Survival Commission (SSC), a branch of theInternational Union for Conservation of Nature and Natural Re-sources (IUCN). The idea is that they should focus on theconservation priorities for a particular group of animals. Byhighlighting problems, debating priorities and suggesting ac-tion, the authors of these plans aspire to stimulate the conserva-

    tion of their group of species. More often than not, the task ofpreparing these plans points more to the breadth of our igno-rance than to the depth of our knowledge, but even that can bestimulating and useful. This booklet, "The Canid Action Plan,"has grown from the deliberations of the IUCN/SSC Canid Spe-cialist Group. The Group was formed in 1981. Fearing the un-wieldiness of large committees, its core membership is smallapproximately one dozen international specialists (see Appen-dix I). However, the Group is supported by an army of affiliatesaround the world, whose efforts have produced the data onwhich this action plan is based (see Apendix 2).

    The Canid Specialist Group has held major meetings inHelsinki (1982), in Edmonton (1985), and in Rome (1989),each time in conjunction with the International TheriologicalCongress. We also met in Krakow, Poland, in 1988 in associa-tion with the International Union of Game Biologists. Thesemeetings have been very fruitful, each introducing a flush ofnew experts to join those stalwarts who have managed to attendthem all. At each, we have benefitted from spoken papers andstimulating discussion. Nonetheless, I think it honest to con-clude that we have felt frustrated that our efforts have led tonothing more substantial than debate.

    To combat this, our first endeavour was to compile a CanidConservation Dossier. This involved the compilation of spe-cies-specific papers which were added to the growing dossier

    ii

  • and circulated quite widely. Notable amongst those earlymembers of the Canid Specialist Group for the effort they putinto the dossier were James Dietz, Lory Frame, Pall Her-steinsson, A.J.T. Jonsingh, James Malcolm,and Patti Moehlman.Our plan had been for the Dossier to grow to a point at which itcould be published as a book. However, that plan was overtakenby events as the top priority became completion of this actionplan, which has, in some respects, usurped some of the roleoriginally envisaged for the Canid Conservation Dossier. Timewill tell how best we should proceed from here, but the impor-tant point is to acknowledge how the efforts of those whoprepared dossier papers contributed to the evolution of thisaction plan.

    In writing this action plan, Joshua Ginsberg and I have, timeand again, been thwarted in our efforts to phrase simple conclu-sions, although we have been vividly aware that, to be effective,the action suggested in such a document should be straightfor-ward. This dilemma is at least partly a reflection of the natureof the Canidae and their diverse dealings with people. TheSimien jackal appears to need seclusion from people and theirdogs, if it is to survive in its highland refuges. The African wilddog's intolerance of human intrusion, and its need for vast spacespell certain conflict with modern man. The bush dog is sorarely seen that we have no idea about its circumstances, butfear the worst. On the other hand, red foxes are notorious fortheir success in urban settings, and plenty of other canids,ranging from coyotes to crab-eating zorros, seem able to thriveamidst human settlements. Management of various canids(largely prompted by rabies control and fur harvest, along withattempts to limit perceived depredation upon domestic stock)leads to the slaughter of hundreds of thousands of foxes annu-ally, thousands of wolves and a handful of African wild dogs.But then, there are many foxes and a fair number of wolves towithstand this onslaught (ill-conceived although it may some-times be), whereas there is scarcely a handful of wild dogs left.When the issues are so disparate, what logical rules are we to usein assigning priorities between the management of the manyand the conservation and management of the few? The logicalmorass thickens with the realization that the fox trotting acrossyour field of view may be, simultaneously, a resource for thetrapper, a health hazard to the rabies official, a quarry to thehuntsman, a subject to the photographer, vermin to the poultryfarmer, and a joy to behold to the aesthete. What is more, theirjudgements are neither right nor wrong according to some self-appointed propheteach could argue a case, but would do sousing incomparable currencies (how are we to equate units ofjobs versus units of cultural heritage versus units of sufferingversus money etc?). Again and again, scientific judgement tripsover ethical judgement.

    In writing this booklet we have tried at least to untangle theseissues, even if we could not resolve them. For me, progress in

    this task took a major bound forward with the arrival in Oxfordof Dr. Joshua Ginsberg. Dr. Ginsberg joined the Wildlife Con-servation Research Unit at Oxford University in 1988, and soonafterwards became Deputy Chairman of the Canid SpecialistGroup. His energy and clarity of thought have had an enormousimpact on this text. He and I would both like to acknowledgeespecially Miss Lynn Clayton, whose groundwork laid thefoundations for our writing. Wolves are watched over by aseparate Specialist Group, and we are grateful to that group and,in particular, its chairman, Dr. David Mech, for supplying theinformation we presenton wolves. Furthermore, we are gratefulfor grants from the International Fur Trade Federation, thePeople's Trust for Endangered Species and from the IUCNthrough the good offices of Dr. Simon Stuart, without whichthis work could not have been done. Preparation of the Plan hasinvolved a mountainous correspondence, the burden of whichfell largely on Dr. Ginsberg's shoulders. He received much ableassistance from Karen da Silva. We list in Appendix 2 thenames and addresses of all those around the world who havecorresponded with us, and we thank them all. In addition, wethank Amie Brutigam, Jeff McNeely, Ulie Seal, Simon Stuartand Rosemary Woodruff for their help with various chapters,and Juliet Clutton-Brock for proofreading the final version.

    Clearly it is my aim, as Chairman of the Group, and co-author of the Plan, to foster sensitive and scientific conservationof the Canidae. The road to this aim involves considerations ofmany turnings: there are the special roles of disease, depreda-tion on stock, and fur harvests in canid conservation; there is thedifficulty of inter-specific competition between similar species,and the perplexing issue of hybridization with domestic dogs.We have raised each of these issues in Chapters 9 to 12. We havebeen unimpressed by some of the cases made against canids,and by the unimaginative nature of many traditional solutions.Yet we have been heartened by some innovative ideas (such asthe oral vaccination of foxes against rabies, and the use ofaversive conditioning to forestall predation). However, beforedeciding which turn to take along the road to conservation ofcanids, one must first ask why travel that road at all. I couldanswer with comments on the stability of communities, thediversity of gene pools, the minimum sizes of viable popula-tions and other more or less abstruse notions. But at the bottomof it, I want canids conserved because I like them. If you likethem too, then hopefully you will find some merit in our recom-mendations. If you do not like them, then my task is to changeyour mind, and because I suspect that this is more likely themore you learn about canids, I am hopeful that this booklet mayspeed your conversion.

    David W. Macdonald, ChairmanIUCN/SSC Canid Specialist Groupand Wildlife Conservation Research Unit

    iii

  • One of the species this action plan covers is the grey wolf (Canislupus). As a species, the wolf still survives in large numbers.However, that is only because it happens to have had the mostextensive original range of any land mammal. The wolf hasbeen exterminated from most of western Europe, the UnitedStates, and Mexico.

    Wolf conservation efforts, then, must be devoted both to bol-stering remnant populations that are threatened or endangered,and to restoring wolves to ecosystems intact enough to supportthem. In the United States, Yellowstone National Park, wherewolves were wiped out over 50 years ago, stands out as a primeexample of the latter. Central Scandinavia represents a similarchallenge in Europe; less than 10 remain there, including onlya single breeding female. The closely related red wolf (Canisrufus) of the southeastern United States is gone from the wildexcept for a few animals recently reintroduced to North Caro-lina.

    Thus this Action Plan proposes such efforts as these, but italso calls attention to the many other wolf populations through-out the northern hemisphere whose status is tenuous or un-known. The human population pressures anticipated during thenext few decades may well threaten several of these poorlyknown wolf populations.

    This makes the Action Plan especially timely. If the wolfrecovery and research efforts proposed herein can be carried outduring the next decade, they probably will make the differencebetween further loss of the wolf and holding the line againstsuch a loss.

    L. David MechChairmanIUCN/SSC Wolf Specialist Group

    Grey wolf (Canis lupus). (Photo by L.D. Mech)

    iv

  • Section 1. Introduction

    1. Objectives, Structure, and Limitationsof the Canid Action Plan

    IntroductionThe Canid Action Plan is one in a series of such plans commis-sioned by the International Union for Conservation of Natureand Natural Resources (IUCN) and written by members of theSpecies Survival Commission's Specialist Groups. The au-thors of action plans have a clear remit: to provide current andaccurate information that will help individuals, institutions, andgovernments to make educated decisions with the aim ofensuring the long-term survival of the species in question. Tosucceed in this objective, the authors of an action plan must firstcollect, collate, and synthesize the information available on thestatus, abundance, and distribution of the taxon under consid-eration. Only then can priorities be established and a plan foraction developed.

    Of course, before proceeding to these higher goals, there aresome basics to be agreed upon. First, which species are to berecognized, and second, by which names are we to refer tothem? These questions are not as trifling as they might at firstappear, because classification of the family Canidae is moder-ately controversial (Wayne et al. 1989), and because there is aprofusion of confusing common and Linnaean names. InChapter 2, we discuss canid taxonomy, and outline the classi-fication that we have adopted here at the species level. Al-though we have not, for the most part, gathered sufficientinformation to analyze conservation priorities at the level ofsubspecies, we have included a list of recognized subspeciesand appropriate citations for those who wish to pursue ques-tions of taxonomy and biogeography.

    Structure of the Canid Action PlanGeographicSome other Specialist Groups have organized the contents oftheir action plans on a regional basis. This makes good sense:for many species, policies are formulated on a regional basis.Hence, a regional approach allows policy makers in differentareas to find, quickly, data which apply to their countries. Someaction plans (e.g. the antelope plans, East 1988) are furtherdivided into regional reports, each of which covers a smallgeographic area, providing a level of detail that is enviable.However, such an approach is not appropriate for the canids,and we have opted to organize the canid action plan differently.The reasons for this are several and stem from the basic biology

    of carnivores: first, predatory species occur at lower densitiesthan prey; second, at any one location, the diversity of carnivorespecies is usually rather low, while third, the geographic rangesof many species of carnivore are rather large. In east andnortheast Africa, for instance, there are 59 species of antelope;on average, each country supports 26 species (East 1988). Incontrast, these 59 species of antelope live side-by side with onlysix species of canid. Each country in the region supports, onaverage, four species.

    These traits of canid distribution are not just regional, butglobal. There are only 34 canid species worldwide. The twocontinents with the highest species diversity, Africa and SouthAmerica, each support a mere 10 species. Most species withinthe Canidae have distributions that span at least a wholecontinent. One species, the red fox, is found throughout theNorthern Hemisphere (and in Australia).

    In deciding how to present our information we were im-pressed by the generalization that the status of a particular canidspecies appears to be remarkably consistent throughout itsrange. Red foxes are common, and thriving, wherever theyoccur; side-striped jackals are rare, but not threatened, through-out their range; the African hunting dog is nearly extirpated inevery country where it is still found. There is a single exceptionwith which to prove the rule: the grey wolf, common in northernCanada and Siberia, is threatened to a point verging on extirpa-tion in much of its range.

    Given the expansive geographical ranges of many canidspecies, and their relatively low species diversity, a country bycountry analysis of status, abundance, and distribution wouldbe unwarranted and repetitive. Furthermore, it would be un-workable insofar as information on the status of canids is veryscattered. For most countries, one simply could not writedetailed reports because detailed information is unavailable.Our generalizations about status are based on the reports of ourcorrespondents, who have reported species as being rare, threat-ened, or abundant. Only in that small minority of cases forwhich detailed surveys have been completed (e.g. the Africanwild dog (Frame and Fanshawe in prep.)) or where an entireSpecialist Group is devoted to a single species (the grey wolf)is the quality of the information remotely satisfactory.

    Nonetheless, policies often have a regional focus. Manypeople involved in biological conservation are shifting theirattention from a species-oriented to an ecosystem-oriented ap-proach. A catalogue of the world's canids, arranged alphabeti-cally, would be cumbersome, and would necessitate a morass ofcross references. Therefore, in what we hope is a sensible and

    1

  • utilitarian compromise between the realities of canid biology,the limitations of the data, and the necessity for a geographicalframework, we have organized the first part of the Canid ActionPlan by geographic regions. The regions we have devisedreflect, in broad terms, the biogeographical distribution ofCanidae. In some cases, following the lead of the canids, wehave crossed continental boundaries. North Africa has beenpaired with the Middle East, Europe with North America.

    Each of the world's canid species has been assigned to oneof the following five chapters. The arrangement of species sec-tions within chapters follows a regional approach:

    Chapter 4. Sub-Saharan Africa: African wild dog; bat-earedfox; black-backed jackal; Cape fox; golden jackal; side-stripedjackal; Simien jackal.

    Chapter 5. South America: Azara's zorro; bush dog; crab-eating zorro; culpeo; grey zorro; hoary zorro; maned wolf;Sechuran zorro; small-eared zorro.

    Chapter 6. Holarctic: Arctic fox; coyote; grey fox; grey wolf;island grey fox; raccoon dog; red fox; red wolf; swift or kit fox.

    Chapter 7. North Africa and the Middle East: Blanford'sfox; fennec; pale fox; Rppell's fox.

    Chapter 8. South and Southeast Asia and Australasia:Bengal fox; corsac fox; dhole; Tibetan fox; dingo.

    An area as large and heterogeneous as "Eurasia and NorthAmerica" may seem excessive for a single chapter. We are notoblivious to the potential absurdity of a single section whichembraces almost all of the Northern Hemisphere. However,only 11 species are found therein, and, of that 11, the distribu-tions of three of the species (the red fox, the grey wolf, and thegrey fox) exceed even the arbitrary boundaries we have set.Similarly, the range of the golden jackal includes much of northAfrica, the Middle East, south and east Asia, and part of Europe.However, most of our information on this species comes fromsub-Saharan Africa, so it seemed appropriate to locate thespecies account in Chapter 4. The structure of each geographi-cally organized chapter is explained in Chapter 3, "Introductionto Geographic Chapters and Species Summaries."

    Structure of the Canid Action PlanGeneralFoxes, wolves, and jackals have much in commonboth intheir biology, and in their relationships with man. Hence,following the geographical chapters, we have written fourchapters on topics almost universally germane to the Canidae asa whole. The topics are trade, captive breeding, predation onstock and game, and disease.

    Perhaps the most frequent contact between many urbandwellers and wild canids is when the latter are encountered inthe form of a fur coat. Clearly the fur trade is a conspicuousaspect of canid management, and although an exhaustive analy-sis of that trade is overdue, and much needed, it would be toolarge an undertaking for this book. Rather, we have focused

    here on the immediate impact of the trade in endangered andthreatened species. Chapter 10 presents an analysis of the tradein canid species listed under the Convention on InternationalTrade in Endangered Species of WildFauna andFlora (CITES).We summarize the absolute levels of trade recorded from 1980-1986, and discuss the patterns of trade and use of canids incommerce.

    For many endangered species, captive breeding and subse-quent reintroduction may offer a final, albeit expensive andcircuitous, form of redemption. Exterminated in the wild, thered wolf (Canis rufus) has recently been bred in captivity andreintroduced in North Carolina (Phillips and Parker 1988).Similarly, kit foxes, extirpated from the northern end of theirrange are now being reintroduced to Canada (Carbyn pers.comm.).

    A successful captive breeding programme is often a pre-requisite for reintroduction. So, in Chapter 11, we analyze datafrom the International Zoo Yearbooks (1963-1984) to discoverthe current level of effort directed at breeding canids in captiv-ity, and the success of these endeavours.

    The very opportunism and mobility that is the hallmark ofsuccess for many members of the dog family is also responsiblefor throwing them into conflict with people. This conflict leadsus to consider two further topics: first, competition betweenman and canids through depredations on game and domesticstock; and second, canids as victims of, and often vectors for,several zoonoses, of which rabies is undeniably the mostnotable. In addition to rabies, there are several other pathogensof direct(e.g. leishmaniasis) and indirect (e.g. sarcoptic mange)concern to man, the latter threatening the fur trade, the formerthreatening life.

    The traditional response to perceived problems of predationand disease has been to attempt to reduce canid numbers bykilling them. This time-honoured approach has two notabledrawbacks: it tends to throw different factions into conflict (e.g.fur traders object to their commodity being blasted by irate shot-gun-toting stockmen, and those concerned with animal welfareobject to both groups killing canids); and it may not work(either because the problem was merely perceived but not real,or because the solution was inappropriate).

    The issues raised by predation and disease are biologicallyand economically complicated. We have tried, in Chapters 12and 13, to expose the salient arguments. In Chapter 12, weconsider aspects of predation by wild canids on livestock andgame. Although many canid species are widely held to have asignificant economic impact on the numbers of valuable herbi-vores, dispassionate analyses of these accusations are few.Because data are often scanty and unevenly spread betweenspecies, our review is biased towards those few canids for whichmore information is available. Our intention is to identifyproblems common to many canids, real and imaginary, to seeksome indication of their magnitudes, and to evaluate the effec-tiveness of existing solutions and the practicability of novelones, all from the point of view of species conservation.

    In Chapter 13 we discuss disease amongst canids, and itsimplications for their conservation and for public health. Dis-ease poses both direct and indirect threats to populations of wildcanids. In the case of direct threats, the burden of our argument

    2

  • is that disease can decimate small populations of canids (andother animals) irrespective of any security they might appar-ently be guaranteed by being protected in reserves and parks.We present this argument by assessing the prevalence ofdisease in various canid populations, along with evidence thatdisease operates as a population regulation mechanism incanids and, hence, has the potential to wipe out "viable"populations.

    Disease is also an indirect threat to the conservation ofcanids. Many canids may act as vectors for pathogens that posea threat to humans (or their livestock and pets). Hence, canidsare frequently killed in attempts to control zoonoses. Ourarguments are based on a review of the evidence that diseasesare harboured in canid populations, consideration of the effectsof "control" programmes on canid populations, and the efficacy(and hence, economic/ecological wisdom) of such "control"measures.

    Notes on Data CollectionThe species accounts presented here were compiled by thefollowing process: census forms were sent to people known tobe interested in canids; these forms posed questions about thedistribution and status of wild canids, and solicited informationon the names and addresses of other authorities whom weshould contact; data garnered through this survey and from areview of published literature were combined in a draft entry foreach species; each such draft was sent to several reviewers;reviewers were selected, where possible, to span the geographicregion covered by the species in question; and corrections madeby reviewers were incorporated in the preparation of a finalversion of each species' entry. The Canid Action Plan could nothave been completed without the help of these correspondents,and we have tried to represent their views accurately. Nonethe-less, we alone are responsible for any errors that may remain inthe text.

    Many of our correspondents have emphasized the potentialusefulness of a register of those involved with canid research.In response, we have included such a list in Appendix 2.Doubtless we have not searched widely enough in compilingthis directory, but if there are embarrassing omissions we canonly hope that this will prompt readers to advise us of them sothat they may be remedied in future editions.

    Limitations of the Canid Action PlanThe structure we have imposed upon this first edition of theCanid Action Plan has suited our purpose, but it has drawbacks.For example, by making our focus regional and global, we havelargely ignored aspects of conservation at the level of sub-species or of local populations. In defense of this, we wouldargue that, for the most part, patterns of extinction are regional:a succession of sub-populations disappears, survivors becomefragmented, and local extirpations start the slippery slide toextinction. In addition, for many species such as the bush dogor the fennec, data are so inadequate that the available materialsswiftly determined the scope of our summary. While we mayhave struggled to float our ideas at a regional level, they wouldundoubtedly have sunk at any finer resolution. However, for aminority of species, such as the wolf and the African wild dog,we have presented a summary of information for each sub-population, made a stab at assigning priorities among the sub-populations, and directed readers to publications that coverthese species in greater detail.

    The nomination of priorities also lures us into an imponder-able mirage dividing biology and ethics. It may be tempting toseek criteria on which to decide whether it is more important tosave the Simien jackal than the red wolf. But both are irre-placeable. We might be able to guess the order in whichendangered canid species will go extinct, but we have tried notto place relative value on the loss of one versus another.Nonetheless, as funds and time are limited, we have tried tofocus our action planning on those species most threatened withimminent extinction.

    Finally, we are mindful that this is the first edition of theCanid Action Plan. We see its publication as a starting point, ajob begun rather than one completed. We hope that it willstimulate a steady stream of information to the Canid SpecialistGroup, and that this information will lead to a subsequentedition in which our lens can focus at a finer level of resolution.Ideally, a future edition will treat every species at the level thathas been possible here only for the wolf and wild dog. In themeantime, while we should plan for the worst, we can hope forthe best. Perhaps the gaps in our information house the reser-voirs of elusive species; perhaps the fennec fox is abundantsomewhere in north Africa, the dhole in Burma, the wild dogsin Ethiopia. Let us hope so, indeed, let us try hard to find out,but let us not bank upon it.

    3

  • 2. Sorting out the Canidae

    The Importance of TaxonomyIn this chapter, we describe the classification of the familyCanidae which we are using for this action plan. Classification,or taxonomy, is all too often viewed as an arcane branch ofbiology. Taxonomy reflects our understanding of evolutionand ecology, and therefore is critical to developing sound con-servation practices and priorities (Wilson 1988a). Taxonomygoes well beyond naming animals; it allows us to make rela-tively objective statements about the relationships within andbetween groups of organisms. It enables us to place boundariesaround variation so that we can make the judgement, forexample, that a fox found in Korea is a subspecies of the red foxand not an entirely new species. Making such distinctions is aprerequisite to planning how to conserve the greatest naturaldiversity.

    Ideally, we would preserve a representative sample of anentire species' genetic diversity. The distinction betweenspecies and subspecies, or even between subspecies and racewould be irrelevant (Ryder 1989). Ideally, sufficient funds,sufficient land, and sufficient interest would exist so that wecould preserve species as the evolutionary units that they are, oronce were. Unfortunately, and invariably, conservation ispractised in a world far removed from this Utopia. Conservationpriorities, both in the field and in zoos, must be judged with apragmatic eye to the allocation of resources and manpower(Conway 1986; Leader-Williams and Albon 1988).

    At the very least, preserving sufficient intra-specific vari-ation to sustain a species is, almost by definition, a prerequisiteto its conservation. The goal of many conservation efforts,often unstated, appears to be to preserve as many species aspossible (see papers in Wilson 1988b). While biologists mayargue the benefits of conserving the greatest intra-specificdiversity or the greatest inter-specific diversity, such an argu-ment assumes that political decisions are made with regard tobiology. Often they are not. Policy makers, constrained byfunds, space, and politics, must often base their actions onpolitical, not ecological, priorities. As biologists, the least wecan do is clearly define what we consider the species, orsubspecies, involved in the equation.

    Laws are often written to protect species, not subspecies.Clearly, the theoretical matter of how species are defined be-comes critical, as does the practical matter of whether a givenpair of populations differ at the species level. Acknowledgingthat the species is the taxonomic unit or level at which manybiological debates are conducted, it becomes crucial to considerwhether it is also the level at which to address legislation. Yetthe fact remains that before we can discuss the conservation

    status of a particular species, or group of species, we must becertain that what we are calling a species is really just that, andnot "merely" a subspecies of something else.

    Classification of SpeciesTable 1 lists the species of Canidae and the names by which werefer to them. The classification follows closely that of Corbetand Hill (1980) (following that of Clutton-Brock et al. 1976)with several exceptions discussed below.

    We hope that this classification will be regarded as a con-structive compromise in the context of conservation strategies.In the 14 years since the publication of Clutton-Brock et al.'spaper, of the 34 species listed, we have adopted only threechanges in specific status. These include two types of changes:two (or more) species listed by Clutton-Brock et al. may havebeen amalgamated into a single species; or specific status maybe granted to a group which Clutton-Brock et al. considered asubspecies of a larger species group. The three changes we havemade are:

    1.

    2.

    3.

    Dusicyon culpaeus/culpaeolus. Following Corbet and Hill(1980), we have included Dusicyon culpaeolus as a subspe-cies of D. culpaeus. The description of D. culpaeolus wasmade from a single skull and skin in the British Museumcollection (Clutton-Brock et al. 1976). The major differencebetween the two species is one of size, and D. culpaeusexhibits large variation in size.

    Dusicyon griseuslfulvipes. D. fulvipes, morphologicallyvery similar to D. griseus (Clutton-Brock et al. 1976), wasuntil recently thought to be restricted to the island of Chiloe.A recent discovery shows, however, that there are mainlandpopulations of this group (Medel et al. 1990). Given thisevidence, there is a strong argument for separating the twospecies. Indeed, we strongly recommend further analysis(genetic and/or morphological) to confirm the specific statusof D. fulvipes. However, in the interests of parsimony, anduntil such analyses are undertaken, we follow the classifica-tion of Corbet and Hill (1980) and group it with D. griseus.

    Vulpes veloxlmacrotis. Corbet and Hill (1980) treat thesetwo as separate species. Recent reviews (O'Farrell 1987,Scott-Brown et al. 1987) follow this classification. How-ever, Hall (1981) suggests that the two are, in fact, a singlespecies. In the southern part of their range, the two aresympatric and hybrids have been recorded in New Mexico(Rohwer and Kilgore 1973) and Texas (Thornton et al.

    4

  • Table 1. Common EnglishAction Plan

    Scientific Name

    Alopex lagopusCanis adustusCanis aureusCanis familiaris dingoCanis latransCanis lupusCanis mesomelasCanis rufusCanis simensisCerdocyon thousChrysocyon brachyurusCuon alpinusDusicyion culpaeusDusicyon griseusDusicyon gymnocercusDusicyon sechuraeDusicyon vetulusDusicyon microtisFennecus zerdaLycaon pictusNyctereutes procyonoidesOtocyon megalotisSpeothos venaticusUrocyon cinereoargenteusUrocyon littoralisVulpes bengalensisVulpes canaVulpes chamaVulpes corsacVulpes ferrilataVulpes pallidaVulpes rueppelliVulpes veloxVulpes vulpes

    names used in the Canid

    Common Name

    Arctic foxSide-striped jackalGolden JackalDingoCoyoteGrey WolfBlack-backed jackalRed wolfSimien JackalCrab-eating zorroManed wolfDholeCulpeoGrey zorroAzara's zorroSechuran zorroHoary zorroSmall-eared zorroFennec foxAfrican wild dogRaccoon dogBat-eared foxBush dogGrey foxIsland grey foxBengal foxBlanford's foxCape foxCorsac foxTibetan foxPale foxRppell's foxSwift, or kit foxRed fox

    1971). Furthermore, recent genetic evidence indicates thatthese two are, in fact, a single species (Dragoo pers. comm.).Given both the genetic and biogeographic evidence, we havefollowed Hall's (1981) classification.

    In addition to changes in specific status, we differ fromCorbet and Hill (1980) in several generic classifications. These,arguably, are less important in their implications for conserva-tion. For example, none of our recommendations, nor anylegislation real or imagined, is likely to be affected whether weallocate the fennec fox to the genus Fennecus or Vulpes. Atleast for the moment, nobody has proposed conservation triagedecisions at the level of the genus. Nonetheless, for the sake ofcompleteness, we have included the results of recent researchby adopting the following changes:

    1. Urocyon species. Corbet and Hill (1980) list both Urocyonspecies as members of the genus Vulpes. However, morpho-logical, karyological, palaeontological, and allozyme evi-dence (Wayne and O'Brien 1987; Collins in press) allsuggest that Urocyon is a distinct genus which has a rela-tively early origin (6-9 million years ago) and that the islandand mainland grey foxes are distinct species. Following thisevidence, we have assigned both species of North Americangrey fox to Urocyon.

    2. Fennecus zerda. Corbet and Hill (1980) refer to the fennecfox as a member of the genus Vulpes on the basis of its

    skeletal and cranial characteristics. Genetic evidence, whileplacing the fennec in the Vulpes group, indicates that genericstatus is probably warranted (Wayne and O'Brien 1987).

    3. Cerdocyon thous. The South American crab-eating fox isfrequently placed in its own genus, Cerdocyon (see Berta1982). However, Corbet and Hill (1980) suggest that thedifferences between the crab-eating fox and other membersof the Dusicyon group are insufficient to warrant a separategenus. Recent genetic studies do little to resolve the genericstatus of this species, beside confirming that it is closelyrelated to members of the Dusicyon group (Wayne andO'Brien 1987). Given the widespread use of the Cerdocyonclassification, and considering the equivocal nature of thegenetic evidence, but acknowledging that the genetic studiestend to indicate some degree of divergence from the Dusicyongroup, we have opted to avoid confusion by using the genusCerdocyon.

    In Table 2, we list 223 subspecies of the world's canids. Thislist is by no means definitive, nor is it meant to be a revision ofthe family. Rather, we hope it will provide a guide to thesubspecific diversity of the Canidae. Further information on thedistribution of each subspecies is given in the primary refer-ences cited in the minority of cases where it is available.

    Common English NamesThis text is written in anticipation of an international reader-ship. This poses a problem in so far as national names are wildlyambiguous. On the other hand, to resort solely to Linnaean, orscientific, names might be considered the hallmark of a turgidtext. In our compromise, we have opted for those commonEnglish and Spanish names that minimize ambiguity. Nonethe-less, the reader should be aware that one person's silver foxrefers to Vulpes vulpes while another's is Vulpes chama. Equally,in our lifetimes, and in British English, Canis simensis has beenknown, successively, as the Abyssinian wolf, the Simien fox,and, most recently, as the Simien jackal. The message, perhapscounterintuitive to the non-specialist, is that it is generallysimpler to work with scientific names.

    The names we use to identify a particular species must berecognizable to all our readers, whether we use the commonname in English or the scientific name. In this context, we alertthe reader to particular confusion in the use of common namesfor the small canids in South America. Frequently, small SouthAmerican canids in the genera Dusicyon and Cerdocyon arecalled foxes, although this term might best be applied to speciesin the genus Vulpes. Furthermore, both Dusicyon culpaeus andVulpes vulpes are commonly called the red fox. Likewise, bothDusicyon griseus and Urocyon cinereoargenteus are com-monly called the grey fox. One cumbrous solution would be tocall these animals the South American grey and red foxes. Oursolution reflects our geographical bias in retaining the word"fox" for the Northern Hemisphere species and referring toDusicyon culpaeus as the culpeo and Dusicyon griseus as thegrey zorro. Hopefully readers will be tolerant of our use ofanglicized Spanish in an attempt to develop English commonnames which will be unambiguously understood.

    5

  • Table 2. Recognized subspecies in the Family

    Scientific Name

    Alopex lagopus beringensisAlopex lagopus fuliginosusAlopex lagopus groenlandicusAlopex lagopus hallensisAlopex lagopus lagopusAlopex lagopus pribilofensisAlopex lagopus sibiricusAlopex lagopus spitzbergensisAlopex lagopus ungavaCanis adustus adustusCanis adustus bwehaCanis adustus centralisCanis adustus kqffensisCanis adustus lateralisCanis aureus algirensisCanis aureus anthusCanis aureus aureusCanis aureus beaCanis aureus lupasterCanis aureus maroccanusCanis aureus ripariusCanis aureus soudanicusCanis familiaris dingoCanis latrans cagottisCanis latrans clepticusCanis latrans dickeyiCanis latrans frustorCanis latrans goldmaniCanis latrans hondurensisCanis latrans impavidusCanis latrans incolatusCanis latrans jamesiCanis latrans latransCanis latrans lestesCanis latrans mearnsiCanis latrans ndcrodonCanis latrans ochropusCanis latrans peninsulaeCanis latrans texensisCanis latrans thamnosCanis latrans umpquensisCanis latrans vigilisCanis lupus albusCanis lupus alcesCanis lupus arabsCanis lupus arctosCanis lupus baileyiCanis lupus bernardiCanis lupus campestrisCanis lupus chancoCanis lupus columbianusCanis lupus crassodonCanis lupus fuscusCanis lupus griseoalbusCanis lupus hudsonicusCanis lupus irremotusCanis lupus labradoriusCanis lupus ligoni

    Canidae

    Taxonomist

    MerriamBechsteinBechsteinMerriamLinnaeusMerriamDybowskiBarret-Hamilton & BonhoteMerriamSundevallHellerSchwarzNeumannScalterWagnerCuvierLinnaeusHellerHemprich & EhrenbergCabreraHemprich & EhrenbergThomasMeyerHamilton-SmithElliotNelsonWoodhouseMerriamGoldmanAllenHallTownsendSayMerriamMerriamMerriamEschscholtzMerriamBaileyJacksonJacksonMerriamKerrGoldmanPocockPocockNelson & GoldmanAndersonDwigubskiGrayGoldmanHallRichardsonBairdGoldmanGoldmanGoldmanGoldman

    Year

    1902179917991900175819021922189818841846191419151902184018411820175819141833192118321903179318391903193218511904193619031934191218231897189718971829189718971949194918971792194119341935192919431804186319411932183918231941193719371937

    Reference

    Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Hall 1981Hall 1981Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Corbett and Newsome 1975Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Hall 1981Hall 1981Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981

    6

  • Table 2. (continued)

    Scientific Name

    Canis lupus lupusCanis lupus tycoonCanis lupus mackenziiCanis lupus nubilusCanis lupus occidentalisCanis lupus orionCanis lupus pallipesCanis lupus pambasileusCanis lupus signatusCanis lupus tundrariumCanis mesomelas achrotesCanis mesomelas arenarumCanis mesomelas elgonaeCanis mesomelas mesomelasCanis mesomelas schmidtiCanis rufus gregoryiCanis rufus rufusCanis simensis citerniiCanis simensis simensisChrysocyon brachyurusCuon alpinus adust usCuon alpinus alpinusCuon alpinus dukhunensisCuon alpinus fumosusCuon alpinus hesperiusCuon alpinus infuscusCuon alpinus javanocusCuon alpinus lanigerCuon alpinus UpturnsCuon alpinus primaevusCuon alpinus sumatrensisDusicyon culpaeus andinusDusicyon culpaeus culpaeolusDusicyon culpaeus culpaeusDusicyon culpaeus lycoidesDusicyon culpaeus magellanicusDusicyon culpaeus reissiiDusicyon culpaeus smithersiDusicyon griseus domeykoanusDusicyon griseus fulvipesDusicyon griseus gracilisDusicyon griseus griseusDusicyon griseus maullinicusDusicyon gymnocercus antiquusDusicyon gymnocercus gymnocercusDusicyon gymnocercus incaDusicyon sechuraeDusicyon vetulusDusicyon microtisCerdocyon thous aquilusCerdocyon thous azaraeCerdocyon thous entrerianusCerdocyon thous germanusCerdocyon thous thousFennecus zerdaLycaon pictus lupinusLycaonpictus manguensisLycaon pictus pictusLycaonpictus sharicus

    Taxonomist

    LinnaeusSchreberAndersonSayRichardsonPocockSykesElliotCabreraMillerThomasThomasHellerSchreberNoackGoldmanBachmande BeauxRuppelIlligerPocockPallasSykesPocockAfanasiev & ZolotareuPocockDesmPocockHeudeHodgsonHardwThomasThomasMolinaPhillippiGrayHilzheimerThomasPhillippiMartinBurmeisterGrayPhillipiAmeghinoFischerThomasThomasLundSclaterBangsWiedBurmeisterAllenLinnaeusZimmermannThomasMatschieTemminckThomas and Wroughton

    Year

    17581775190818231829193518311905190719121926192619141778189719371851192218351811194118111831193619351936182019361892183318211914191417821896183619061914190118371861183619031889181419141914184218821898182418611923175817801902191518201907

    Reference

    Ellerman and Morrison-Scott 1966Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Hall 1981Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Hall 1981Hall 1981Coetzee 1977Coetzee 1977Cabrera 1961Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Chasen 1940Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Chasen 1940Cabrera 1961Corbet & Hill 1980Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Corbet & Hill 1980Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Corbet & Hill 1980Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Cabrera 1961Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977

    7

  • Table 2. (continued)

    Scientific Name

    Lycaon pictus somalicusNyctereutes procyonoides koreensisNyctereutes procyonoides orestesNyctereutes procyonoides procyonoidesNyctereutes procyonoides ussuriensisNyctereutes procyonoides viverrinusOtocyon megalotis megalotisOtocyon megalotis virgatusSpeothos venaticus venaticusSpeothos venaticus wingeiUrocyon cinereoargenteus borealisUrocyon cinereoargenteus californicusUrocyon cinereoargenteus cinereoargenteusUrocyon cinereoargenteus colimensisUrocyon cinereoargenteus costaricensisUrocyon cinereoargenteus floridanusUrocyon cinereoargenteus fraterculusUrocyon cinereoargenteus furvusUrocyon cinereoargenteus guatemalaeUrocyon cinereoargenteus madrensisUrocyon cinereoargenteus nigrirostrisUrocyon cinereoargenteus ocythousUrocyon cinereoargenteus orinomusUrocyon cinereoargenteus peninsular isUrocyon cinereoargenteus scottiiUrocyon cinereoargenteus townsendiUrocyon cinereoargenteus venezuelaeUrocyon littoralis catalinaeUrocyon littoralis clementaeUrocyon littoralis dickeyiUrocyon littoralis littoralisUrocyon littoralis sanatacruzaeUrocyon littoralis santarosaeVulpes bengalensisVulpes canaVulpes chamaVulpes corsac corsacVulpes corsac kalmykorumVulpes corsac turkmenicaVulpes ferrilataVulpes pallida edwardsiVulpes pallida hartertiVulpes pallida oertzeniVulpes pallida pallidaVulpes rueppelli caesiaVulpes rueppelli cufranaVulpes rueppelli rueppelliVulpes rueppelli somaliaeVulpes velox arsipusVulpes velox deviaVulpes velox hebesVulpes velox macrotisVulpes velox muticaVulpes velox neomexicanaVulpes velox nevadensisVulpes velox tenuirostrisVulpes velox veloxVulpes velox zinseriVulpes vulpes abietorum

    Taxonomist

    ThomasMoriThomasGrayMatschieTemminckDesmarestMillerLundIheringMerriamMearnsSchreberGoldmanGoodwinRhoadsElliotAllen and BarbourMillerBurt and HopperLichvensteinBangsGoldmanHueyMerriamMerriamAllenMerriamMerriamGrinnell and LindsdaleBairdMerriamGrinnell and LindsdaleShawBlanfordSmithLinnOgnevOgnevHodgsonRochebruneThomas and HintonMatschieCretzschmarThomas and Hintonde BeauxSchinzThomasElliotNelson and GoldmanMerriamMerriamMerriamMerriamGoldmanNelson and GoldmanSayBensonMerriam

    Year

    19041922192318341907184418211909183919111903189717751938193818951896?18991941185018991938192818911899191119031903193018581903193018001877183317581935193518421883192119101826192119391825191819041909190218881902190219311931182319381900

    Reference

    Coetzee 1977Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Coetzee 1977Coetzee 1977Cabrera 1961Cabrera 1961Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Cabrera 1961Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Hall 1981Ellerman and Morrison-Scott 1966Ellerman & Morrison-Scott 1966Coetzee 1977Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman & Morrison-Scott 1966Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Coetzee 1977Hall 1981Hall 1981Hall 1981Hall 1981HaU 1981Hall 1981HaU 1981Hall 1981Hall 1981Hall 1981Hall 1981

    8

  • Table 2. (continued)

    Scientific Name

    Vulpes vulpes aeygptiacaVulpes vulpes alascensisVulpes vulpes alpherakyiVulpes vulpes alticolaVulpes vulpes anatolicaVulpes vulpes arabicaVulpes vulpes atlanticaVulpes vulpes barbarasVulpes vulpes beringianaVulpes vulpes cascadensisVulpes vulpes caucasicaVulpes vulpes crucigeraVulpes vulpes dauricaVulpes vulpes dilutaVulpes vulpes dolichocraniaVulpes vulpes dor sailsVulpes vulpes flavescensVulpes vulpes fulvaVulpes vulpes griffithiVulpes vulpes harrimaniVulpes vulpes hooleVulpes vulpes ichnusaeVulpes vulpes indutaVulpes vulpes jakutensisVulpes vulpes japonicaVulpes vulpes karaganVulpes vulpes kenaiensisVulpes vulpes krimeamontanaVulpes vulpes kurdistanicaVulpes vulpes macrouraVulpes vulpes montanaVulpes vulpes necatorVulpes vulpes ochroxanthaVulpes vulpes palaestinaVulpes vulpes peculiosaVulpes vulpes pusillaVulpes vulpes regalisVulpes vulpes rubricosaVulpes vulpes schrenckiVulpes vulpes silaceaVulpes vulpes splendidissimaVulpes vulpes stepensisVulpes vulpes tobolicaVulpes vulpes tschiliensisVulpes vulpes vulpes

    Taxonomist

    SonniniMerriamSatuninOgnevThomasThomasWagnerShawMiddendorffMerriamDinnikBechsteinOgnevOgnevOgnevGrayGrayDesmarestBlythMerriamSwinhoeMillerMillerOgnevGrayErxlebenMerriamBraunerSatuninBairdPearsonMerriamOgnevThomasKishidaBlythMerriamBangsKishidaMillerKishidaBraunerOgnevMatschieLinnaeus

    Year

    181619001906192619201902184318001875185219141789193119241926183818431820185419001870190719071923186817771900191419061900183619001926192019241854190018971924190719241914192619071758

    Reference

    Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Hall 1981Hall 1981Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966Ellerman and Morrison-Scott 1966

    9

  • Section 2. Species and Geographic Analyses

    3. Introduction to Species Summariesand Geographical Chapters

    Format of Regional ChaptersTo facilitate access to information, much of the data on thestatus, abundance, and distribution of the world's 34 canidspecies are organized in Chapters 4-8. Each of these chaptersis organized in a similar format.

    In the following section, we have reproduced the headingsused in each of the geographic chapters. Under each heading,we discuss the information included in that part of the chapter.

    IntroductionThe introduction to each chapter includes a general discussionof the geographic area included therein, the number of speciesin the chapter, and which species occur in the region covered.If a species occurs in the region, but is included in a differentchapter, this is cross-referenced.

    Current Status of SpeciesA brief summary of the current conservation status of eachspecies is presented in this part of each chapter. Full details ofthe conservation status of all the canid species, and recom-mended conservation actions, can be found in Chapter 13.

    Species AccountsFollowing the summary sections at the beginning of eachchapter, we include an entry for each species found in thatparticular geographic region. For each species, an account hasbeen prepared which uses the following format. Data areincluded for each variable where available.

    Name. English, scientific,French, German, local names (whereknown). Where more than one English name is commonly used,we include the preferred name in the title, and list others in thissection. Local names are also presented.

    Description. General physical description for identificationincluding: head-and-body length, tail length, shoulder height,weight.

    Reproduction. Data on time of mating, gestation, litter size,period of lactation, age at sexual maturity, and longevity.

    Social Behaviour. Information concerning general patterns ofactivity, mating system, parental behaviour, patterns of dispersal.

    Diet. Range of diet and feeding preferences.

    Distribution. Geographical range of the species. For rarespecies, or species about which little is known, specific loca-tions may be included.

    Habitat. Ecological range of the species.

    Population and Status. Known patterns of distribution andabundance. For common species, information is general andnotes the species' ability to coexist with man outside protectedareas are included. For threatened species, detailed statusreports have been included. Single sightings of rare or unknownspecies may be included if data are few.

    Commercial Use. Details of commercial uses, both locally andin terms of international trade.

    Other Threats. Threats to the species other than trade.

    Current Research Programmes. Details of research pro-grammes known to be active listing objectives and principal in-vestigators.

    Proposed Research Programmes. Proposals for further re-search.

    Conservation Measures Taken. Occurrence of viable popu-lations in National Parks and reserves. For common species, upto five protected areas providing viable populations are listed.Details of specific legal protection for threatened species aregiven. For rare species, all reserves in which a species is knownto occur are listed. General legal status in each country (restric-tions on internal trade, hunting/trapping regulations, etc.) willnot usually be included.

    Reviewers. Names of scientists who have reviewed our draftversions of this section.

    10

  • 4. Sub-Saharan Africa

    IntroductionSub-Saharan Africa covers a vast area which encompassesexamples of nearly every tropical and sub-tropical biome.Nonetheless, only seven species of canid are found in this partof the continent. Five of these species are found only in sub-Saharan Africa: the side-striped jackal (Canis adustus); theblack-backed jackal (Canis mesomelas); the Simien jackal(Canis simensis); the bat-eared fox (Otocyon megalotis); andthe Cape fox (Vulpes chama). The African hunting dog (Lycaonpictus) was formerly found in parts of north and northwestAfrica, but is now extirpated in these regions. The golden jackal(Canis aureus) is still found in many areas of the Middle andNear East and southern Europe. However, because much of theinformation available for the species is derived from researchconducted in sub-Saharan Africa, we have summarized the in-formation available on the golden jackal in this section.

    Side-striped jackal (Canis adustus). (Photo by J.R. Malcolm)

    Current Status of SpeciesThe overall status of canids in sub-Saharan Africa is good. Fiveof seven species are abundant and face no serious threats. Twospecies, the wild dog and the Simien jackal, however, are indanger of extinction. Whatever the status of a particularspecies, however, our knowledge of the behaviour and ecologyof most species which occur in sub-Saharan African is good,surpassed as a group only by the general knowledge we haveabout species occurring in Eurasia and North America. Further-more, there are active population monitoring efforts in manyparts of Africa. Finally, in much of Africa, there is greatconcern, both local and international, about the fate of wildlife.Although concern does not always translate into protection, itis unlikely that an extinction could occur without sufficientwarning to address the causes of decline.

    Species AccountsDetailed biological studies of each species are listed in thereference section following this chapter. Although there hasbeen no thorough review of African canids, several textsprovide basic information about the distribution, abundance,and biology of African mammals. These include Haltenorth

    and Diller (1980); Kingdon (1977); Smithers (1983); and Stuart(1981).

    Side-striped jackal (Canis adustus)

    French: Le Chacal flancs rays. German: Streifenschakal.Afrikaans: Witwasjakkals. Karamojong: Oloo. Kikinga: Ngwe.Kinyakyusa: Akambwe, Imbira. Kinyiha: Habila. Kiswahili:Bweha, Bweha miraba. Luganda: Akabowa, Ekihe. Lugbara:Bowa. Lwo: Too. Madi: Uba. Runyankole: Emuha. Runyoro:Eboha. Sebei: Bleyit.

    Description. A greyish yellow coat with a white stripe fromelbow to hip with a white tip on the tail (may be absent). Stripesnot always obvious. Head-and-body length: 65-81 cm. Taillength: 30-41 cm. Shoulder height: 41-50 cm. Weight: 6.5-14kg, males somewhat larger than females, males mean of 9.4 kg,females 8.3 kg.

    Reproduction. Time of mating: Jun.-Jul./Sept.-Oct. (E. Af-rica); Jun-Nov. (S. Africa). Gestation: 57-70 days. Litter size:3-6, mean 5.4. Lactation: 8-10 weeks. Age at sexual maturity:6-8 months, disperse at 11 months. Longevity: 10-12 years.

    Social Behaviour. Mated pair and their young, young disperseat 11 months before the next litter. Nocturnal. Not averse tomoving in close proximity to dwellings and have been sighted,like the red fox, in city centres.

    11

  • Diet. Omnivorous, taking a variety of invertebrates, small ver-tebrates, carrion, and plant material. Data from animals takenin agricultural areas of northern Botswana (Smithers and Wilson1979, in Smithers 1983) show a predominantly vegetable diet,supplemented by rodents and insects. Included cow and goathair suggest scavenging of domestic livestock found as carrion.

    Habitat. Avoids open savanna, preferring thickly woodedareas but not forest. Sea level up to 2700 m in East Africa.

    Distribution. Tropical Africa, moist woodlands, 15N to 23S,not in the rain forests of west and central Africa. Prefers moisterparts of savannas, thickets, forest edge, cultivated areas; up to2700 m. West African range is largely unknown but includesnorthern Nigeria (Gombe, north of Benue and Borgu GameReserve in the west). Occurs in the Central African Republic,south Sudan, and parts of Ethiopia, central and southern Kenya,Uganda, and along Lake Victoria. Widespread in Tanzania andZambia, northern Mozambique, Malawi, south, west, and eastZaire and into Gabon. Also Angola, northern Namibia, north-ern Botswana, throughout Zimbabwe, and in the Transvaalsouthwards into Kwazulu and Natal provinces. See Figure 1.

    Figure 1. Distribution of the side-striped jackal (Canis adustus).

    Population and Status. Rare throughout its range, but nodirect threat is known.

    Commercial Use. None known, although uses in traditionalmedicine are known. The heart is used to treat epilepsy in theBuganda tribe. Skins and nails are used to ward off evil spirits.

    Threats. Jackals have historically (1950s) been trapped andpoisoned during rabies outbreaks in Uganda. They are alsothought to have died in large numbers during early parts of thecentury due to distemper.

    Current Research Programmes

    1. Studies are being conducted on diet in the Akagera NationalPark, Rwanda, by J. Kalpers.

    2. Long-term research on the black-backed and golden jackalsin the Serengeti by P.D. Moehlman has been expanded toinclude the side-striped jackal.

    Conservation Measures Taken. Occurs in numerous nationalparks and reserves including: Serengeti National Park, Tanza-nia; Akagera National Park, Rwanda

    Reviewers. P. Moehlman; D. Rowe-Rowe.

    Golden jackal (Canis aureus). (Photo by A.J.T. Johnsingh)

    Golden or Asiatic jackal (Canis aureus)

    French: Le Chacal commun. German: Goldschakal. Kiswahili:Bweha wa mbuga, Bweha Dhahabu.

    Description. Coat varies with season and region, but is usuallya pale gold-brown, or brown tipped yellow. Fur is coarse, andnot very long. Dorsal area is black and grey; head, ears, andsides can be rufous. Underside is frequently ginger or nearlywhite. The tip of the tail is black. There is sexual dimorphismof approximately 15% in body weight. Head-and-body length:60-106cm. Tail length: 20-30 cm. Shoulder height: 38-50 cm.Weight: 7-15 kg.

    Reproduction. Time of mating: Oct.-Nov. (Tanzania occa-sionally June-July, Moehlman pers. comm.); Feb.-Mar. (USSR);Oct.-Feb. (Israel). Timed so birth occurs at peak of food abun-dance (e.g. Thomson's gazelle fawns in the Serengeti, Jan.-Feb.). Gestation: 63 days. Litter size: 1-9, usually 5-6. Lacta-tion: 8-10 weeks. Age at sexual maturity: 11 months (female);

    12

  • up to 2 years (male) but may delay reproduction. Longevity: 16years in captivity (Yom-Tov pers. comm.), up to 13 years in thewild (Moehlman pers. comm.).

    Social Behaviour. Usually in mated pairs, territorial, territorysize 0.5 to 2.5 km2. Helpers (last year's young) can increase re-productive success, but not as much as in the black-backedjackal (Moehlman 1983). In many areas, C. aureus is nocturnaland diurnal, but it becomes strictly nocturnal in areas inhabitedby humans. Deviation from usual social organization is foundin areas where food is abundant, with up to 20 members in agroup and with home range areas of 0.11 km2 (Macdonald1979).

    Diet. Omnivorous: fruits; invertebrates; reptiles; amphibia;birds; small mammals; carrion. Opportunistic foragers, butcooperation between mated pairs can greatly increase huntingsuccess. In areas around human habitation, it can subsist on gar-bage (Macdonald 1979).

    Habitat. Open country with trees and brush, grass and copse.Occur up to 2,000 m (though recently sighted at nearly twicethat altitude). Also found in oases and human settlements in theIsraeli desert. Habitat can vary immensely, including areasaround human habitation.

    Distribution. Widely distributed in north and east Africa,southeast Europe, south Asia to Burma and Thailand. In Africa,there is some evidence of the golden jackal extending its rangein recent times, having lately been sighted in the Bale Moun-tains National Park, Ethiopia, up into the heather belt (3,800 m),also in the south in the Harenna forest.(J.C. Hillman, pers.comm.). Recently sighted in eastern Italy (L. Boitani pers.comm.). See Figure 2.

    Population and Status. Locally abundant and widespread.Accurate estimates of numbers rare, but surveys in the Serengeti(May 1986) estimated approximately 1,600 individuals, withno significant change in the last decade (Campbell and Borner1986).

    Commercial Use. None known.

    Threats. None known.

    Current Research Programmes. Long-term research in theSerengeti, Tanzania, by Dr. P.D. Moehlman.

    Conservation Measures Taken. Occurs in numerous Na-tional Parks and reserves in Africa and Asia.

    Reviewers. P. Moehlman; Y. Yom-Tov.

    Black-backed jackal (Canis mesomelas)

    French: Le Chacal chabraque. German: Schabrackenschakal.Afrikaans: Rooijakkals. Ateso: Ekwee. Karamojong: Kwee.Kigogo: Nhyewe. Kihehe: Nchewe. Kikinga: Ngewe. Kilian-gulu: Gedala. Kinyaturu: Mola. Kisagara: Kewe. Kiswahili:Bweha Nyakundu, Bweha Shaba. Kitaita: Muzozo.

    Description. A reddish brown or ginger canid with a distinc-tive black saddle running from the nape of the neck to the tail.The tail is reddish-brown and black and bushy. Head-and-bodylength: 45-90 cm., means: 72 cm males, 67.3 cm females (Zim-babweSmithers 1983); 78.5 cm males, 74.5 cm females(Cape Province, Stuart 1981). Tail length: 26-40 cm., means:32.9 cm males, 31.4 cm females (Zimbabwe, Smithers 1983);32.6 cm males, 31.6 cm females (Cape Province, Stuart 1981).

    Figure 2. Distribution of the golden jackal (Canis aureus).

    13

  • Black-backed jackal (Canis mesomelas). (Photo by J. Nel)

    Shoulder height: 30-48 cm., mean 38 cm (Zimbabwe, Smithers1983). Weight: 6-12 kg, means: 7.9 kg males, 6.6 kg females(all classes, Zimbabwe, Smithers 1983); 8.4 kg males, 7.4 kgfemales (adults, Natal, Rowe-Rowe 1978); 8.15 kg males, 7.4kg females (adults, Cape Province, Stuart 1981).

    Reproduction. Time of mating: May-Aug. May-Jun. (Natal,Rowe-Rowe 1978); July (Botswana, Smithers 1971). Gesta-tion: 60 days. Litter size: 1 to 8, mean4, 1-3 survive. Lactation:8-10 weeks. Age at sexual maturity: 11 months, but may delayreproduction (Moehlman 1979,1983). Longevity: 14 years incaptivity, but probably usually at most 7 years in the wild(Rowe-Rowe 1982).

    Social Behaviour. Monogamous pairs which mate for life(Moehlman 1983). Often hunt in pairs. Diurnal and crepuscu-lar (see Ferguson et al. 1988 for details); may become noctur-nal in areas around human habitation and in some conservationareas (Kalahari, Kruger National Park). Home range size 2 to 33km2, but larger sizes are rare (references in Moehlman 1986;Ferguson et al. 1983). Rowe-Rowe (1982) found that matedadults had home ranges of 19 km2, young (< 1 year) 9 km2, andunmated adults 33 km2. Helpers (last year's young) maycontribute to rearing of pups and greatly increase pup survival(Moehlman 1979,1983). Sometimes old termite mound oraardvark holes are used as dens, and are usually moved amongseveral dens.

    Diet. Omnivorous and varied. In South Africa, Rowe-Rowe(1983) reports that in a study in Giant's Castle National Re-serve, 55% of the diet was from small mammals, 11 % ante-lopes; 9% hare-sized mammals; 9% birds; and 16% miscellane-

    ous including domestic animals, invertebrates, amphibians,grubs, and fruit. In the Serengeti, a similar diet is taken, withvariation in the amount of mammalian prey being a function ofseasonal abundance (Lamprecht 1978). Moehlman (1979)reports a preference for rodents and the fruit of Balanitesaegyptiaca, which may serve as a natural worming agent.Stuart (1976) found that in the Namib Desert, fruit constituteda large proportion of the diet. Some food may be cached(Lamprecht 1978).

    Habitat. Variable, ranging from human habitation and suburbsof large cities to the desert of the Namib. Found in areas withup to 2,000 mm of rain per annum (Rowe-Rowe 1976), tend tobe replaced by the side-striped jackal in moister areas. Commonin areas with less than 100 mm per annum.

    Distribution. Widespread in east and southern Africa. Twopopulations (not connected) exist. In the north, it is found fromthe Gulf of Aden south to Tanzania. In the south, the populationranges from the Cape north to southwestern Angola and east toZimbabwe and Mozambique. See Figure 3.

    Population and Status. Population density figures are avail-able for only a few sites. In the plains areas of the SerengetiNational Park, Tanzania the species may have exhibited adecline in population from 1976 to 1986 (488 to 50 individuals(Cambell and Borner 1986) with densities as low as oneindividual per 100 km2. However, C. mesomelas populationsare recovering in those areas surveyed. Furthermore, thissurvey did not include many transects in habitat used by C.mesomelas, and hence may underestimate the real populationsize (Moehlman pers. comm.). Rowe-Rowe (1982) estimatesthat jackal densities in the Giant's Castle Game Reserve areapproximately one per 2.5 km2.

    Figure 3. Distribution of the black-backed jackal (Canis mesomelas).

    14

  • Commercial Use. Fur exploited. Trapping, hunting withhounds, and the use of "coyote getter" toxic collars on sheep arelegal in South Africa. In South Africa, it may be huntedthroughout the year.

    Threats. In South Africa, and elsewhere in Africa, the black-backed jackal has a reputation as a voracious killer of lambs andsheep, a reputation that may, in part, be deserved. However, asthe identification of carcasses killed by jackals and other canidsis often not undertaken, deaths caused by domestic dogs may beattributed to wild canids. Predation on sheep is seasonal, andoccurs most frequently during lambing (Rowe-Rowe 1986). InSouth Africa, control measures include trapping and hunting.Extirpation is difficult, however, and the nature conservationdepartment suggests killing only those jackals causing damage(Rowe-Rowe 1986).

    Current Research Programmes

    Simien jackal (Canis simensis)

    English: Simien fox, Abyssinian wolf, red jackal, Ethiopianwolf. French: Loup asyssinie. Amharic: Ky Kebero. Oromo:Jedalla Farda.

    Description. Bright reddish (coat lighter in juveniles andfemales) with white patches on throat, neck and chest, and basalhalf of tail white underside. Lower half of tail black. Relativelylong legs and a long, thin muzzle. Head-and-body length: 100cm. Tail length: 33 cm. Shoulder height: 60 cm. Weight: ap-proximately 15-18 kg males, 13-16 kg females.

    Reproduction. Time of mating: Aug-Dec (Sillero-Zubiri andGottelli unpubl. data) Gestation: 60+3 days. Litter size: 2-6(N=7, Sillero-Zubiri and Gottelli unpubl. data). Lactation: 6-8 weeks. Age at sexual maturity: 2 years, females. Longevity:unknown.

    Social Behaviour. Most frequently seen alone while foragingby day. Pairs and groups of 2-12 congregate at morning, noonand evening with noisy greetings. Adult animals have beenobserved with sub-adults (9 months) sleeping together as a

    Simien jackal (Canis simensis). (Photo by D. Gottelli)

    group. Several adult-sized animals were seen together at a densite that contained pups (Malcolm 1988). The largest groupsighted consisted of 7 adults and 6 pups. Mean group size is 7,often multi-male groups. Fifteen dens have been observed:dens usually consist of a much-used system of burrows beneatha rock-overhang or cliffs; other burrows located in a flat, grassyarea have several entrances, possibly interconnected. Dens areattended both by parents and sub-adult helpers. All members ofthe pack regurgitate food to the young. All information is fromSillero-Zubiri and Gottelli (1989).

    Diet. Subsists primarily on mammals ranging in size from haresand the giant mole-rat Tachyoryctes macrocephalus (900g) tothe common species of grass rat (Arvicanthis blicki, Otomystypus) at 90-120g. Also includes Lophuromys melanonyx,Stenocephalemys albocaudata, and Lepus starcki in diet(Malcolm 1982; Sillero-Zubiri and Gottelli unpublished). TheSimien jackal is a key predator in the food chain of this highaltitude ecosystem, and its role in regulating the dynamics ofrodent populations is important. In Bale, one jackal was seenfeeding off a cow carcass. Caching prey and scavengedmaterial in shallow holes is common (Sillero-Zubiri and Got-telli unpublished). The species has been observed huntingreedbuck and mountain nyala calves. Evidence of other preyhas not been found in scat analysis (Hillman 1986).

    Habitat. Common in open moorlands where vegetation is lessthan 0.25 m high. Also seen in heather moorlands.

    Distribution. Mountains of Ethiopia at altitude range 3,000-4,377 m. Arssi and Bale mountains of southeast Ethiopia; theSimien mountains, northeast Shoa, Gojjam (1932 sighting) andMt. Guna (Yalden et al. 1980). See Figure 4.

    15

    1.

    2.

    3.

    Long-term research being conducted by P.D. Moehlman inthe Serengeti National Park, Tanzania. No other activeresearch projects known.

    Investigations on improving ways of assessing damage tolivestock, preventing damage, and improving methods byselectively eliminating stock-killers are being carried out byresearchers from the University of Natal under the supervi-sion of M.R. Perrins and D.T. Rowe-Rowe.

    C. and T. Stuart of the African Carnivore Survey are in-volved in a project to survey the effect of predator controlmeasures on all carnivore populations in southern Africa.

    Conservation Measures Taken. Occurs in numerous pro-tected areas, including the Serengeti National Park, Tanzania,and the Kruger National Park and Giant's Castle Game Reserve,South Africa.

    Reviewers. G. Mills; P. Moehlman; D. Rowe-Rowe;C. and T. Stuart.

  • Figure 4. Distribution of the Simien jackal (Canis sinensis).

    Population and Status. Total population under 1,000 indi-viduals. The Bale Mountains National Park supports the onlyknown population of any size, estimated at 500-600 individuals(Sillero-Zubiri and Gottelli 1989; Malcolm 1988 estimated450-600). Within the Park, the distribution is as follows:eastern area (Sanetti and Shaiya) 200-250 individuals; westernarea (Web Valley and Peaks) 150-200; other areas 50-100 indi-viduals (Malcolm 1988 as modified by data from Sillero-Zubiriand Gottelli). The maximum number of individuals in the Parkis estimated at 704 (Hillman 1986). Population in Bale is stable(Malcolm 1988) or perhaps increasing (Sillero-Zubiri andGottelli unpublished). In the Simien Mountains, the totalpopulation was estimated to be less than 20 individuals in 1976.This area has been closed for security reasons for some time,and the current status of the population there is unknown(Malcolm pers. comm.). In Arsi, there are no more than 100individuals, and the population is decreasing (Malcolm 1982).In northeast Shoa, recent sightings suggest a small scatteredpopulation (Sillero-Zubiri and Gottelli pers. comm.).

    Commercial Use. None known.

    Threats

    4.

    5.

    6.

    Simien jackal (Gottelli pers. comm.). Overgrazing mayaffect food available for rodents, the jackal's primary food(Malcolm 1988). There is some evidence that some types ofhuman activity (grazing) may increase the habitat availableto the jackals (Sillero-Zubiri and Gottelli unpublished data).

    In the northern parts of their range, jackals have historicallybeen hunted as pests. Evidence exists that they are respon-sible for lamb predation (Sillero-Zubiri and Gottelli: one ob-servation of predation on domestic stock), but this has onlybeen observed within protected areas.

    Genetic introgression by cross-breeding with domestic dogsthreatens to pollute the Bale population gene pool.

    Competition with the golden jackal may, in the future,present a problem, although there is no evidence at presentof interspecific competition.

    16

    1.

    2.

    3.

    Loss of suitable habitat.

    The existence of just three small populations immediatelysuggests that disease is an equally great threat to the species.A single outbreak of distemper, parvo, and/or anthrax coulddecimate the last remaining large population in the BaleMountains. Potential vectors for disease include domesticdogs (C. familiaris).In the Web Valley of Bale, two forms of human-inducedcompetition also threaten the species. Domestic dogs havebeen observed hunting rats and may compete with the

    Current Research Programmes. A study has been funded byWildlife Conservation International (begun in March 1988) onthe ecology, behaviour, and conservation of this species and iscurrently being conducted by C. Sillero-Zubiri and D. Gottelli.

    Conservation Measures Taken. The Bale and Simien popu-lations are completely protected by inclusion in National Parks.The species is protected in Ethiopia by law (Wildlife Conserva-tion Regulations 1974: Schedule VI). The law states that thespecies "may only be hunted with special permit for scientificpurposesonly to be issued by the Minister of Agriculture."No poaching, hunting and/or trade of live animals has been ob-served.

    Reviewers. D. Gottelli; J. C. Hillman; J. Malcolm; C. Sillero-Zubiri. We would like to thank D. Gottelli and C. Sillero-Zubirifor their generous contribution of unpublished data.

  • African wild dog (Lycaon pictus)

    English: Cape hunting dog. French: Lycaon, Le cynhyne,Loup-peint German: Hynenhund. Afrikaans: Wildehond.Ateso: Apeete. Kalenjin: Suyo. Kibena: Liduma. Kibungu:Eminze. Kichagga: Kite kya negereni. Kihehe: Ligwami.Kijita: Omusege. Kikamba: Nzui. Kikuyu: Muthige. Kikuyu,Kimeru: Mbawa. Kiliangulu: Eeyeyi. Kimaragoli: Imbwa.Kinyaturu: Mbughi. Kinyiha: Inpumpi. Kinyiramba: Mulula.Kisukuma: Mhuge. Kiswahili: Mbwa mwitu. Kitaita: Kikwau.Kizigua: Mauzi. Lwo: Sudhe, Prude. Masai: Osuyiani. Sam-buru: Suyian. Sebei: Kulwe, Suyondet.

    Description. Unmistakable colour patterns with splotches ofyellow, black, white and grey covering the entire body. Someregional variation in colours, but intraspecific variation is largein all places. Large rounded ears, long legs, broad bushy tailswith tips usually, but not always, white. Head-and-body length:76-112 cm. Tail length: 30-41 cm. Shoulder height: 61-78 cm.Weight: 17-36 kg (monomorphicmean 25 kg), somewhatlarger in the south.

    Reproduction. Time of mating: Very variable. In the Ser-engeti, mating starts in November, with 75% of litters bornbetween January and June. In Kruger National Park, matingoccurs in April-May. Gestation: 69-73 days. Littersize:2 to l9,average 7 to 10. Lactation: 10 weeks. Age at sexual maturity:females disperse at approximately 18 months (Frame andFrame 1976), but due to suppression of breeding in all butdominant female and male, actual age at first reproductionusually much greater.

    Social Behaviour. African wild dogs live in packs composedof several related adult males, and one to several related adultfemales originating from a different pack. Pups remain in theden for two to three weeks, then emerge to receive food fromreturning pack members. The wild dog exhibits a complexsocial organization in which all members of the pack feedyoung and mother at den. The number of adults and yearlingsin each pack varies locally, with a mean of 6.6 in the Serengeti(Malcolm and Marten 1982), 8.8 in the Zambesi Valley (Childes

    African wild dog (Lycaon pictus). (Photo by J. R. Ginsberg)

    Figure 5. Distribution of the African wild dog (Lycaon pictus) 1983after Smithers 1983.

    1985), 14.8 in Hwange National Park, Zimbabwe (Childes1985), and 7.3 in Kruger National Park (Reich 1981). Totalpack size, including pups, can be quite variable and will dependon pack fecundity and pup survival. The species is perhaps oneof the few truly nomadic animals. Pack home range variesconsiderably, ranging from 500 km2 at its smallest (Reich 1981)up to 1500 km2 (Frame et al. 1979). Pack home range overlapvaries from 50% to 80%. In most circumstances, in any particu-lar pack, only the dominant female will rear pups successfully.The dominant bitch will remain at the den with the pups and befed, along with the pups, by other members of the pack whoregurgitate food after returning from a kill.

    Diet. Variable according to prey species most abundant in area.Dominant species include: impala in Kruger National Park(Reich 1981); duiker and reedbuck in Zambia (Shenton and Uysin Smithers 1983); Thomson's gazelle and wildebeest in theSerengeti (Schaller 1972; Frame 1986); impala and kudu inHwange National Park, Zimbabwe (Childes 1985).

    Habitat. Very catholic in its requirements, being found in areasof moderately dense bush, open plains habitat, and up into thelower forests of Mt. Kenya.

    Distribution. African wild dogs are never even locally abun-dant, and their nomadic movements make even simple esti-mates of distribution difficult. The population status for eachcountry is listed below. As the status of this species appears tobe changing rapidly, we have included two distribution maps(Figures 5 and 6). Until very recently (Smithers 1983; Figure 5)the range of the wild dog included much of Africa, as far westas Ivory Coast, southern Nigeria, Burkina Faso, around LakeChad, and in western Chad. Disjunct populations formerlyoccurred in southern and southwestern Algeria, the Tanzerouftand Adrrar des Forar in Air in Niger, and near Tibesti in extreme

    17

  • Figure 6. Distribution of the African wild dog (Lycaon pictus)datafrom Frame and Fanshawe in prep.

    north Chad. The species ranged from the Central AfricanRepublic, into southern Sudan, across southern Ethiopia andinto Somalia. They were found throughout Kenya, Tanzania,eastern Rwanda, southeastern and eastern Zaire, Zambia,Malawi, Mozambique, Zimbabwe, Botswana, and Angola.The southernmost populations were found in Namibia, Botswanaand northern South Africa. Although formerly widely distrib-uted in South Africa, by 1983 the wild dog was only found inKruger National Park.

    The wild dog is one of the few canids for which good data areavailable on the historical contraction of the species range. Thespecies ranged over much of what is now South Africa, occur-ring at relatively high densities compared to those now ob-served. The species probably occurred as far south as the Capein 1684 (Smithers 1983). In Zimbabwe, the distribution is nowlimited to areas in and around national parks, reserves, andhunting areas, having been exterminated from all other areas.

    One must remember that even in 1983, within the boundariesof this vast area, the distribution of the wild dog was verypatchy. Since then, the distribution of the wild dog has con-tracted rapidly. Within the boundaries delineated in the mapabove, perhaps 2000 wild dogs can be found. Figure 6 is agraphic depiction of the data presented in the next section.

    Population and Status. Frame and Fanshawe (in prep.) are inthe process of summarizing the data collected from a continent-wide mail survey investigating the status of the wild dog. Avery brief summary of their findings follows:

    Algeria: suspected to be extirpated.

    Angola: very rare, near extirpation.

    Benin: suspected to be extirpated.

    Botswana: still present and moderately abundant for Lycaon.However, despite legal protection, can be shot if a threat tolivestock.

    Burkina Faso: present, but very rare, in southern part of thecountry.

    Burundi: suspected to be extirpated or nearly so.

    Cameroon: small numbers and populations stable. Possi-bility for recovery.

    Central African Republic: rare, but present. Not protectedin any way.

    Chad: Lake Chad and the west. Little known of popula-tions.Congo: suspected to be extirpated.

    Djibouti: probably extirpated.

    Ethiopia: present but rare in most areas. Relatively com-mon in the southwest, but rare nonetheless.

    Gabon: suspected to be extirpated.

    Ghana: suspected to be extirpated or will soon be so.

    Ivory Coast: very rare.

    Kenya: present and in decline. Much interest in conserva-tion in this country, but increasing human population sizeand livestock numbers are a threat.

    Malawi: present only in national parks. Viable populationonly in Kasungu National Park.

    Mali: very rare; suspected to be extirpated.Mozambique: endangered and soon to be extirpated.Namibia: declining population due to hunting/shooting asvermin. Population perhaps viable, but under threat (C. andT. Stuart, pers. comm.).

    Niger: if present, only in south of country. Soon to beextinct.

    Nigeria: nearly extinct.

    Rwanda: suspected to be extinct.

    Senegal: rare, but present in several locations. Populationsthreatened.

    Somalia: present in south and northeast, but probably de-clining.

    South Africa: endangered (R.S.A. Red Data Book) anderadicated from much of the country. Still viable in KrugerNational Park. Despite the size of the park, however, indi-viduals stray. One farmer bordering the park is known tohave shot 20 dogs during 1988 (G. Mills, pers. comm.).Another radio-collared individual was shot in Mozambique.Population introduced to Umfolozi/Hluhluwe complex, Na-tal.

    Sudan: present in the south, rare to common. Civil war,hence no knowledge of population trends.

    Tanzania: present in many parks and reserves, populationssmall and fluctuating (18 to 100 in the Serengeti). Legallyprotected.

    Togo: suspected to be extirpated.

    18

  • Uganda: extirpated, except for occasional visitors fromTanzania or Sudan (Olivier pers. comm.).

    Zaire: suspected to be extirpated.

    Zambia: present in many areas, but declining. Have beenaffected by recent outbreak of anthrax in the Luanga Valley(Saigawa pers. comm.).

    Zimbabwe: rare and declining, but may be increasing inHwange National Park. Recently afforded legal protection.Strong opposition to conservation measures by both tradi-tional and game ranchers.

    2.

    3.

    4.

    5.

    6.

    7.

    was re-established by J. Fanshawe and Dr. C. FitzGibbon ofthe Serengeti Wildlife Research Institute, full-time monitor-ing has been continuing. Work is currently executed by K.Laurenson and Dr. M. Borner of the Serengeti EcologicalMonitoring Programme. This work involves keeping a pho-tographic file of each individual in the population, radio-collaring and blood sampling dogs for disease and geneticanalysis, and monthly radio-tracking flights to locate packson a regular basis.

    The Kruger National Park is home to another long-termstudy which was concentrated around the work of A. Reich(Reich, 1981). The study is now coordinated by Dr. G. Mills.Three levels of research are underway: Dr. Mills is conduct-ing a study of predator-prey relationships in the park and apack of dogs has been included in this; a photographic file isbeing established with help from the public (in collaborationwith another researcher, A. Maddock); the use of satellitetransmitters is being developed in association with Dr. M.Gorman. This last aspect should allow monitoring the packs,especially if they leave the park and come into conflict withfanners.

    Researchers have begun to assess population trends through-out Botswana, but work has concentrated in the north, withinChobe and Ngamiland Districts, an area of approximately120,000 km2, which includes the protected areas of MoremiNational Reserve, Chobe National Park, Nxai Pan NationalPark and Makgadikgadi Game Reserve. Data are beinggathered by a combination of tactics that include extensivesurveying of people (tourists, guides) entering the studyarea, and the radio-collaring and tracking of packs. A keyaim is to assess the nature of wild dog interactions withlivestock. All of this should lead to conservation recommen-dations for wild dogs throughout Botswana. The fieldresearch director is J. Bulger and the project director is W. J.Hamilton.

    In Kenya, P. Kat, National Museum of Kenya, Nairobi, hasbegun a study which aims to assess the genetic structure ofseveral packs in Kenya. The project, at present, does notinclude an ecological component. Genetic analyses onspecimens collected here, and elsewhere in east and south-ern Africa are being analyzed by R. Wayne. J. Fanshawe andP. Kat are also continuing long-term monitoring of otherpacks of wild dogs throughout Kenya.

    A project in Hwange National Park, and the adjacent huntingand forestry areas, has been started by J.R. Ginsberg, OxfordWildlife Conservation Research Unit. The aims of thisproject are to assess the genetic structure of the populationand screen for disease and parasites, study the ecologicalbasis of nomadism and its implications for conservation, andinvestigate the conflicts (disease, crop raiding) betweenhuman and wild dog populations. The project is funded bythe National Geographic Society and the American Philo-sophical Society.

    Monitoring of hunting, trade, and population trends by theAfrican Carnivore Survey, C. and T. Stuart.

    A project to reintroduce the wild dog to the area aroundMkomazi, Tanzania, is being studied by T. FitzJohn.

    19

    In their summary, Frame and Fanshawe (in prep.) have notestimated population sizes for any of the countries in whichLycaon is found. They note that the species