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Status Survey and Conservation

Action Plan

Wild Cats
Compiled and edited by
Kristin Nowell and Peter Jackson
IUCNESCNJCN
The World Conservation

n

Union

Chicago Zoological

SPECIES

SoaetySURVIVALCat Specialist GroupCAT

COMMISSIONWWFCONSEKVATIONSPECIALISTINTERNATLONALGROUP

IUCN/Species Survival Commission
Conservation Communications Fund andContributors to WiZd CatsThe National Wildlife Federation (NWF) makes a significant annual contribution to the SSC Conservation
Communications Fund, in addition to grants for in situ
conservation coordinated by the SSC. NWF is the largest
non-governmental, non-profit conservation-education andadvocacy organization in the United States. It emphasizes
assisting individuals and organizations of all cultures, inthe United States and abroad, to conserve wildlife andother natural resources and to protect the earth’s environment to assure a peaceful, equitable, and sustainable future.

In 1992, IUCN’s Species Survival Commission established the Conservation Communications Fund to garnersupport for its expansive Publications Programme which
promotes conservation by: (1) providing objective scientific information about biodiversity, habitats, and ecosystems; (2) identifying high priority actions for conservation;and (3) delivering the information and recommendations
to natural resource managers, decision-makers, and others
whose actions affect the conservation of biodiversity.
The SSC’s Action Plans (Wild Cats is #28 in the series),
Occasional Papers, newsletter (Species), membershipdirectory, and other publications are supported by a wide
variety of generous donors (see below):

The World Wide Fund for Nature (WWF) provides significant annual operating support to the SSC. WWF’s contribution supports the SSC’s minimal infrastructure andhelps ensure that the voluntary network and Publications
Programme are adequately supported. WWF aims to conserve nature and ecological processes by: (1) preserving
genetic, species, and ecosystem diversity; (2) ensuring thatthe use of renewable natural resources is sustainable both
now and in the longer term; and (3) promoting actions to
reduce pollution and the wasteful exploitation and consumption of resources and energy. WWF is one of theworld’s largest independent conservation organizations,with a network of national organizations and associates
around the world and over 5.2 million regular supporters.
WWF continues to be known as World Wildlife Fund in
Canada and in the United States of America.

The Sultanate of Oman established the Peter Scott
IUCN/SSC Action Plan Fund in 1990. The Fund supports
Action Plan development and implementation; to date,more than 80 grants have been made from the Fund toSpecialist Groups. As a result, the Action Plan Programme
has progressed at an accelerated level and the network has
grown and matured significantly. The SSC is grateful to
the Sultanate of Oman for its confidence in and support for
species conservation worldwide.
The Chicago Zoological Society (CZS) provides significant in-kind and cash support to the SSC, including grants
for special projects, editorial and design services, staff secondments, and related support services. The president of
CZS and director of Brookfield Zoo, George B. Rabb,
serves as the volunteer Chair of the SSC. The mission of
CZS is to help people develop a sustainable and harmonious relationship with nature. Brookfield Zoo carries out
its mission by informing and inspiring 2 million visitors
annually, by serving as a refuge for species threatened withextinction, by developing scientific approaches to manage species successfully in zoos and the wild, and byworking with other zoos, agencies, and protected areas
around the world to conserve habitats and wildlife.

Publication of Wild Cats was made possible withgenerous grants from WWF-Netherlands
and WWFInternational.
Other contributors include Conservation International andthe International Fur Trade Federation.

ii

Contents
Page

Page

Foreword ......................................................................vii
Elizabeth Marshall Thomas

2. North Africa and Southwest Asia.. ...................... .36
Vulnerability Index to Speciesof the Region (Box 1).......................................... 36
Asiatic lion, Panthera leo persica............................ 37
Cheetah,Acinonyx jubatus ....................................... 41
Leopard,Panthera pardus ....................................... 44
Sandcat, Felis margarita ......................................... 47
Caracal, Caracal caracal ......................................... 50

Acknowledgements .......................................................ix
Introduction ..................................................................xi
Executive Summary ....................................................xii
Global Ranking of Cat Species...
Vulnerability (Box 1) .........................................x111
RegionalRanking of Cat Species
Vulnerability (Box 2) .........................................xiv

3. Tropical Asia ........................................................... 54
Vulnerability Index to Speciesof the Region (Box 1).......................................... 54
Tiger, Panthera tigris ............................................... 55
Borneanbay cat, Catopumabadia........................... 65
Clouded leopard,Neofelis nebulosa....................... .66
Asiatic golden cat, Catopumatemmincki............... .69
Flat-headedcat, Prionailurus planiceps................. .70
Rusty-spottedcat, Prionailurus rubiginosus.......... .72
Fishing cat, Prionailurus viverrinus ....................... .74
Marbled cat, Pardofelis marmorata........................ .76
Leopard, Panthera pardus ....................................... 78
Jungle cat, Felis chaus ............................................. 83
Leopard cat, Prionailurus bengalensis................... .85
Iriomote cat, Prionailurus bengalensis
iriomotensis......................................................... 88

Taxonomy of the Felidae ...........................................xvi
Classificationof the Felidae (Box 1)
W. Christopher Wozencraft ...................................xvii
The History of Felid Classification...
Lars Werdelin........................................................xv111
Molecular Geneticsand Phylogeneticsof the Felidae...
StephenJ. O’Brien ................................*............... xx111

Part I
SpeciesAccounts

4. Eurasia ..................................................................... 90
Vulnerability Index to Species
90of the Region (Box 1)

Introduction ................................................................... 1
Structure of the SpeciesAccounts ............................. 1
Categorization of SpeciesVulnerability ................... .2
Worksheet Summary for Global Cat Species
Vulnerability Rankings(Box 1)............................ 3
1994IUCN ThreatenedSpecies
Categories(Box 2)................................................. 5..........................................

Asia Sub-region ...................................................... 91
Snow leopard, Uncia uncia ...................................... 91
Chinesemountaincat, Felis bieti ............................. 96
Manul, Otocolobusmanul........................................ 97
Asiatic wildcat, Felis silvestris,ornata group ........ .99
Eurasianlynx, Lynx lynx ........................................ 101

I Sub-Saharan Africa ................................................. 7
Vulnerability Index to Speciesof the Region (Box 1)............................................ 7
Black-footed cat, Felis nigripes................................. 8
African golden cat, Profelis aurata ........................ .10
Cheetah,Acinonyx jubatus ....................................... 12
African lion, Panthera leo........................................ 17
Serval, Leptailurus serval ........................................ 21
Leopard,Panthera pardus ....................................... 24
Caracal,Caracal caracal ......................................... 30
African wildcat, Felis silvestris,libyca group ........ .32

Europe Sub-region ............................................... 106
Iberian lynx, Lynx pardinus ................................... 106
Europeanwildcat, Felis silvestris,
silvestris group................................................... 110
5. The Americas ........................................................ 114
Vulnerability Index to Speciesof the Region (Box 1)........................................ 114
Kodkod, Oncifelis guigna ...................................... 115
Andean mountain cat, Oreailurusjacobitus.. ....... .116
Jaguar,Panthera onca............................................ 118
Oncilla, Leopardustigrinus ................................... 122

.*.
III

Page

Page

Mar-gay,Leoparduswiedi ...................................... 124
Canadalynx, Lynx canadensis............................... 126
Geoffroy’s cat, Oncijelis geoflroyi ........................ I29
Puma,Puma concolor ............................................ 131
The Florida Panther(Box 2) ............................. 135
Ocelot, Leoparduspardalis.................................... 137
Bobcat, Lynx rufis .................................................. 140
Pampascat, Oncifelis colocolo .............................. 144
Jaguarundi,Herpailurus yaguarondi .................... .146

Laboratory-basedResearch.................................... 209
Intraspecific Diversity and Systematics:........................ .209
The Questionof Subspecies
SubspeciesIdentification Incorporating
Molecular Genetics(Box 1) ........................ .2 10
Genetics............................................................. 211
PopulationViability Analysis ........................... 213
Infection and Disease........................................ 216
Summary and Conclusions.................................... 218
4. Trade ...................................................................... 220
Introduction ............................................................ 220
The Convention on International Trade inEndangeredSpeciesof Wild Faunaand
Flora (CITES) (Box 1) ...................................... 221
International Trade in Cat Furs .............................. 223
The Biological Impact of Trade on
Wild Populations............................................... 227
SustainableUseof Cats for the Fur Trade:
The North American Example .......................... 228
Review of Bobcat and CanadaLynx
ManagementProgramsin the UnitedStatesand Canada......................................... 228
Habitat Evaluation and Management.......... .230
Assessmentsof Population Size,
Structure, and Trends .............................. 230
Harvest Control andMonitoring ................. .232
Are CanadaLynx and Bobcat Harvests
SustainableasPresently Managed?............ .233
DoesCornrnercialUseBenefit Bobcat andCanadaLynx Conservation?....................... .235
Illegal Trade in Cat Products................................. 236
Illegal Trade in Peltsand Live Animals .......... .237
The Bonesof a Dilemma:
Tigers and Oriental Medicine ..................... ,239
Summary and Conclusions.................................... 243

Part II
Major Issuesin Cat Conservation
1. Cats and Habitat Loss.......................................... 149
Introduction ............................................................ 149
Habitatsfor Cats..................................................... 149
Habitat Classificationand Species
Associations(Box 1) ......................................... 150
Habitat Lossand Fragmentation:
An Overview of Global Trends........................ .151
Habitat Loss.................................................. 151
Habitat Fragmentation.................................. 166
Implications for Cat Species.................................. 169
The Role of ProtectedAreas in Cat
Conservationand the Need for Linkages......... .172
Summary and Conclusions.................................... 179
2. Management of Big Cats Near People.............. .I80
Introduction ............................................................ 180
Cat Predationon Livestock .................................... 180
Managementto Minimize the Problemof
Livestock Lossesto Big Cats............................ 183
ProblemAnimal Control .............................. 184
PredatorsandFarmers(Box 1) ................... .185
Improving GeneralAnti-predator
Livestock Management................................ 185
Compensationfor Livestock Losses........... .186
ProgramsWhich Make Wild Landsan Economically Competitive Form of Land Use......... 188
Tourism and Trophy Hunting ..................... .189
Summary and Conclusions.................................... 191
Big Cat Attacks on People(Box 2). ...................... .192

5. Cats in Captivity ................................................... 244
Introduction ............................................................ 244
A Brief History of Catsin Captivity ..................... .244
Statusof Captive Populations................................ 246
Reproduction in Captivity ...................................... 248
SpeciesBred in Captivity .................................. 248
Captive Breeding and Population
ManagementPrograms................................ 254
A Global Captive Action Plan for Felids..........254
Advances in AssistedReproduction................ .258
Linking Ex situ and In situ Cat Conservation........259
Maintaining Viable Captive Populationsof Rare Species............................................. 260
Research............................................................ 261

3. Research ................................................................ 196
Introduction ............................................................ 196
Field Studies........................................................... 198
Natural History .................................................. 198
Population StatusSurveys................................. 202
Long-term Studies............................................. 206
Resolving Conflicts with People...................... .209

iv

Page

Public Education................................................261
Helping to Pay the Costsof
Wildlife Conservation..................................262
6. Reintroduction ......................................................263
Introduction ............................................................263
Reintroductions......................................................263
Translocationsand Population
Supplementation................................................265
Summaryof Problemswith ReintroducingCats....268
ReleaseAreas Acceptable to People.................268
ReleaseAreas with Adequate andSuitableFood Supply ...................................268
Sourceof Animals to be Reintroduced.............268
DiseaseRisks.....................................................269
Costsof Reintroduction.....................................269
Conclusions:What is the Role of Reintroduction
in Cat Conservation7. ....................................*....270

Color Plates...............................................after page 148
Wild Cats of Africa ..............................................Plate 1
African lion, North African lion (characteristics),
African leopard,African golden cat
Wild Cats of Africa ..............................................Plate 2
African cheetah,Saharancheetah,
King cheetah,Serval
Wild Cats of Africa ..............................................Plate 3
Caracal,Black-footed cat, African wildcat,
SandcatWild Cats of the Americas ..................................Plate 4
Jaguar,Puma,Jaguarundi,Pampascat
Wild Cats of the Americas ..................................Plate 5
Ocelot, Margay, OncillaPart III
An Action Plan for Cat Conservationin the 1990s.................................................................271
Introduction ............................................................271
List of Priority Projects..........................................271
Priority Projectsfor Cat Conservationin the 1990s........................................................275
I. GeneralTopics-Projects 1- 17 ...............275
II. SpeciesProjects-Projects 18-105........280

Wild Cats of the Americas ..................................Plate 6
Canadalynx, Bobcat, Geoffroy’s cat, Kodkod,
Andean mountain cat

Appendices .................................................................307
1. ClassicallyDescribedCat Subspecies...................307

Wild Cats of Asia ..................................................Plate 8
Asiatic cheetah,Asiatic lion, Amur leopard,
Cloudedleopard,Marbled cat

2. Scientific Postmortem:A Protocol forCollection of Data and Specimens
Andrew Kitchener, StephenMcOrist, andRobert K. Wayne....................................................314
3. Scientific Namesof SpeciesMentioned
in the Text ...............................................................317
4. Species-HabitatAssociations...........*..................... 319

Wild Cats of Asia ..................................................Plate 7
Bengal Tiger, Amur (Siberian) tiger,
Caspiantiger, South China tiger,
Indo-Chinesetiger, Bali tiger, Javan tiger,
Sumatrantiger

Wild Cats of Asia ..................................................Plate 9
Asiatic goldencat, Asiatic golden cat (spotted),
Bomeanbay cat, Bomeanbay cat (painting)
Wild Cats of Asia .......**...*................................... Plate 10
Leopard cat, Iriomote cat, Fishing cat,
Flat-headedcat, Rusty-spottedcat, Junglecat

5. Cat SpecialistGroup Members..............................326
6. A Statementby the International Fur
Trade Federation.....................................................331

Wild Cats of Eurasia ..........................................Plate 11
Snow leopard,Eurasianlynx, Iberian lynx,
Chinesemountain cat

7. List of Maps, Figures, and Tables..........................332
References..................................................................335

Wild Cats of Eurasia ................*......................... Plate 12
Europeanwildcat, Asiatic wildcat, ManulFore wordin the attics of houses set in rice fields, and that African
golden cats, which have never been studied but are known
to inhabit rain forests, may include as prey small primates
who fall from the trees and lie injured on the ground. Suchobservations, anecdotal and fragmented though they may
be, are nevertheless precious for two reasons: first, theirvery existence makes it clear that if we don’t preserve thespecies, these passing observations may provide the only
knowledge we’ll ever have; and second, as a whiff of salt
air suggests the ocean: tiny bits of information, however
incomplete, suggest an entire lifestyle, in all its complexity, of animals that at this point we know little about, andthat the vast majority of us will surely never see.
The mass of data assembled by the authors is analyzed
to present general principles of conservation giving a clear
sense of where the priorities for future conservation lie.
These are summarized at the end of each of the Major
Issues chapters. The Action Plan itself (Part III) translates the principles into concrete action-theseprojects
should be carried out in the 1990s to improve the conservation of vulnerable cat species.
But the most important contribution that a book on any
animal can make is to the future of its subject. Time is getting short for many of the cats discussed here. In particular, the big cats- tigers, lions, leopards, jaguars, snow
leopards, and cheetahs-face
the hostility of farmers
because of real and perceived threats to livestock and,sometimes, people. These cats are often killed indiscriminately and their future outside well-protected areas is inserious jeopardy. In one of the most important chaptersof this book, Kristin Nowell and Peter Jackson review thesituation and discuss measures to minimize livestock predation so that big cats outside protected areas can co-exist
with people. This is of vital importance because mostreserves are far too small to accommodate viable big cat
populations with a good long-term chance of survival.
A new and grave threat, with ancient roots, is the hunting of tigers and other big cats for bones for traditional
medicine in China and elsewhere in Asia. This is causinga marked decline in tiger numbers, and in late 1992, Peter
Jackson predicted that, unless current trends were sharply
reversed, the tiger faced virtual extinction in the wild withina decade. Since then it is estimated that, in India alone, over
600 tigers have been poached, while, in Russia, Siberian
tigers have been reduced from around 300 to fewer than200. Large numbers of contraband skins and bones havebeen seized, but they can only be the tip of the iceberg.
Wild cats should not be seen merely as beautiful, but
of little practical value. The cats are part of the web of life,

It is indeed an honor to write a foreword for a book of thisstature-a more comprehensive work than this is hard to
imagine, and a more welcome addition to the store of information on the cat family would be impossible to find. With
this work the authors have set a new standard of scholarship for studies of the cat family. The level of scholarshippresented here, as this work clearly shows, is quite obviously nothing less than the finest and most meticulous.
The aim of the authors is a high one-keepinga tradition of learning that began with compiling data upon thevarious genera of plants and animals of the different continents (I recall a massive multi-volume work entitled The
Lemons and Limes of Siam), a tradition that came into its
own in 1964 with the publication of Ernest P. Walker’s
Mammals of the World. The authors have included within
a single work the entire spectrum of factual literature on
the biology, ecology, distribution, and conservation statusof each member of the cat family, presented in summary
form, providing a comprehensive overview of these fascinating animals so that conservationists
now and in the
future will have a ready reference. Whether a reader islooking for bibliography on a species, the names by whicha certain cat is known in local dialects, the use to which acheetah puts his dew claws, the impact of the fur trade on
the Brazilian jaguar population, or the likely impact of new
Spanish highways on the Iberian lynx population, they willfind what they seek in these pages.
The 1,500 references included here comprise a literature that begins with Marco Polo in the 13th century andextends to the present, and that must represent no less than
5,000 scholar/years of collective effort by various authors.
Until the completion of Wild Cats such a literature could
only have been found by combing libraries throughout theworld. Conservationists in far corners of the world lack
the resources to make those searches. But now, the authorsof Wild Cats have done it for them.
Only a deep and abiding dedication to the cat tribe
could possibly inspire such a magnificent effort, so to an
aficionado such as myself, this book is mouthwatering.
Here it is, you say to yourself as you open the pages withreverent anticipation, Here it all is. And sure enough, one
need only to let the book fall open for something to spring
off the page-from the (to me) grim report that the trophy
hunting of lions by sportsmen brings twice as muchincome to a certain African country as can be derived fromthe viewing of lions by tourists in the national parks, tothe perils awaiting those who would attempt to identify
individual mountain lions by their tracks, to the tantalizing fact that rusty-spotted cats sometimes keep their kittens

viiForeword

the mutual interaction of animals and plants, which underpins human life on Planet Earth.
By its very existence this marvellous work seems toecho Walker’s words in Mammals of the World, to whom
the great biologist dedicated his enterprise. “To the mammals, great and small,” he wrote, “who contribute so muchto the welfare and happiness of man, another mammal,

but receive so little in return, except blame, abuse, andextermination.”
Here, in the hands of Kristin Nowell, Peter Jackson, andthe IUCN/SSC Cat Specialist Group, the cat family is atlast well-served.
Elizabeth Marshall

..s

VIII

ThomasAcknowledgements

Acknowledgements
Wild Cats: Status Survey and Conservation Action Plan
is the result of the generous help of numerous individuals
and institutions named below. To all of them we offer oursincere gratitude.
The Research Librarian of the Cat Specialist Group,
Gail Foreman, laid the groundwork for Wild Cats: Status
Survey and Conservation Action Plan by conducting a literature review and producing detailed information sheetson the country-by-country
conservation status of most of
the wild cat species. Other data sheets were written by Urs
Breitenmoser, Ravi Chellam, Rodney Jackson, A.J.T.
Johnsingh, Kurt A. Johnson, Gary M. Koehler, Daniel
Kraus, Laurie Marker-Kraus, and Chris Stuart. Jill Mellen
was a major contributor to the chapter on cats in captivity.
Kurt Johnson was a major contributor to the trade chapter. Colin Groves and Alan Shoemaker helped shape thelist of classically described felid species which is includedin Appendix 1.
The Etosha Ecological Institute, Etosha National Park,
Namibia, and the Species Survival Commission office at
IUCN Headquarters, Gland, Switzerland, provided officefacilities for Kristin Nowell.
World Wide Fund for
Nature-International
supported the Action Plan-related
activities of Peter Jackson.
Stephen Nash and Kathy Odell, whose work was supported by Conservation International, produced cameraready art from roughly-drawn cat distribution maps.
The World Conservation Monitoring Centre’ s Wildlife
Trade Monitoring Unit (and especially John Caldwell) andTRAFFIC International (especially Teresa Mulliken)
provided detailed trade data and advice on interpretation ofthat data. Several of TRAFFIC’s regional and national
offices reviewed the trade chapter and contributed information for it. Charles Dauphine of the Canadian Wildlife
Service and Peter Meszaros of Statistics Canada provided
harvest and price data for the Canada lynx. Officials fromthe U.S. Fish and Wildlife Service provided useful comments on the management of bobcats for the trade chapter.
WCMC’s Protected Areas Data Unit made their files
available for review (thanks to Graham Drucker).
WCMC’s Habitats Unit (Corinna Ravilious and Richard
Luxmoore) contributed the habitat maps which appear inPart II Chapter 1. Julia Watts of the U.S. Department ofEnergy’s Oak Ridge National Laboratory provided key
data on habitat change in the 1980s.
Staff of the Species Survival Commission (Simon
Stuart, Mariano Gimenez-Dixon, Linette Humphrey, Tim
Sullivan, Susan Tressler, Diane Cavalieri, Karin Nelson,
Gabriella Allen, Clotilde Mack, and Doreen Zivkovic)
provided support throughout for preparation of this volume.

Comments on various drafts came from (in alphabetical order): Ablimit Abdukadir (China); Eve Abe (Uganda);
Marcellin Agnanga (Congo); Ashiq Ahmad (management);
Mikhail Akhverdian (Armenia); Anada Tiega (Niger andnorthwest Africa); Penny Andrews (cats in captivity);
Simon Anstey (Angola and Liberia); Marcel0 Aranda(Mexico, research); Marc Artois (disease); Conrad Aveling(Congo); Juliette Bailey (trade); Theodore Bailey (leopard, Canada lynx); Richard Barnes (leopard); Vandepitte
Bart (Botswana); Klaus Berkmiiller (Laos); Hu Berry(Namibia); Brian Bertram (lion, research); R.S. Bhadauria(caracal in India); Sylvain Biquand (Saudi Arabia); Allard
Blom (Zaire); Ashish Bodasing (trade); J. du P. Bothma(South Africa); Stan Boutin (Canada lynx); Tony and Mary
Jane Bowland (South Africa); Nanette Bragin (ocelot in
captivity); Urs and Christine Breitenmoser (Eurasian andCanada lynxes, management); Steven Broad (trade);
Warren Brockelman (Thailand); Dan Brooks (Paraguay);
Alexander Bukhnicashvili(Georgia); Arturo Canedi(Argentina);
Tim Caro (cheetah); Mateus Chambal(Mozambique);Philippe Chardonnet (West Africa);
Divyabhanusinh Chavda (India); Ravi Chellam (Asiaticlion, management); Peter Christie (cats in captivity); Karen
Corbett (fishing cat in captivity); Ian Coulson (Zimbabwe);
Peter Crawshaw(Brazil, management); Ralph Daly
(Oman); Samantak Das (India); Glyn Davies (Kenya);
Miguel Delibes (Iberian lynx); Teruo Doi (Iriomote cat);
Alain Dragesco-Joffe (Niger); Betsy Dresser (cats in captivity); Holly Dublin (Kenya); Nigel Dunstone (Peru,
research); Sarah Durant (cheetah); Barbara Durrant(clouded leopard in captivity); R. Eagan (Canada lynx,
trade); John Eisenberg (Latin America); Louise Emmons(Latin America); Alexander Esipov (Central Asian republics); Robert Evans (black-footed cat in captivity); N.
Fairall (South Africa); Pier Lorenzo Florio (trade); Gail
Foreman (cat species); Joseph Fox (snow leopard andEurasian lynx); George Frame (cheetah); Helen Freeman(snow leopard); Liza Gadsby (Nigeria); Gao Xingyi(China); Bill Gasaway (research); John Gasperetti (Saudi
Arabia); Aadje Geertsema (serval); Gonzalo Gonzales(Chile); Ute Grimm (trade); Paule Gros (cheetah); Colin
Groves (taxonomy); Juan Carlos Guix (Brazil); Ha Dinh
Due (Vietnam); Elke Hahn (trade); Stephen Halloy (Andean mountain cat); Kevin Hansen (puma); Pave1 Hell(Czech republic and Slovakia); Veronique Herrenschmidt(management); Osvaldo Nestor Herrera (kodkod); Jesse
Hillman (Eritrea and Ethiopia); Rafael and Almeira
Hoogesteijn (Venezuela, management); Bernard HoppeDominik (Ivory Coast); Maurice Homocker (puma); F.C.
Hurst (Nigeria); International Fur Trade Federation (trade);Acknowledgements(lion); Ulysses Seal (Florida panther); John Seidensticker(tiger, management); Assad Serhal (Lebanon); Gary Sharp(cheetah); Alan Shoemaker (leopard, cats in captivity);
Albert0 Simonetta (Somalia); Alex Sliwa (black-footedcat); Brian Slough (Canada lynx, trade); Koen de Smet
(Algeria and northwest Africa); J.L. David Smith (fishingcat, tiger, research); Philippe Stahl (European wildcat, disease); Philip Stander (Namibia); Chris Stuart (southern
Africa, research); Simon Stuart (issues in cat conservation); Mel Sunquist (Bornean bay cat, tiger, Latin
America); Wendell Swank (jaguar); Tan Bangjie (China);
Jose Lobao Tello (Central African Republic); Valmik
Thapar (tiger, India); Jay Tischendorf (U.S.A.); Arlen
Todd (Canada lynx); Schwann Tunhikorn(tiger,Thailand); Siima Umar (Turkey); Chris Vaughan (Costa
Rica); Juan Villalba-Macias(Latin America, trade);
Jacques Verschuren (Central Africa); John Visser (blackfooted cat); Clive Walker (South Africa); Kamal Wassif(Egypt); Carlos Weber (Chile); Lars Werdelin (Canada
lynx, taxonomy); Robert Wiese (cats in captivity); David
Wildt (assisted reproduction); Won Pyong-Oh (Korean
peninsula); Michael Woodford (disease); Sejal Worah(rusty-spotted cat); Anne Wright (India); Alfred0 Ximenez(Brazil); D.W. Yalden (Ethiopia); Shigeki Yasuma(Borneo, Iriomote cat); Jinping Yu (leopard cat); James
Zacharias (rusty-spotted cat).
Christine Breitenmoser-Wiirsten,
Dan Cao-Sheng,
Wynand du Plessis, Cecile Thiery, and Adrienne andPaddy Jackson translated documents from Afrikaans,
Chinese, French, German, Italian, Russian, and Spanish.
Our spouses, Tom Preisser and Adrienne Jackson,
helped in innumerable ways.
The Cat Specialist Group expresses its gratitude to thefollowing persons who have donated illustrations: J.J.
Aldama, Tom Brakefield,
Ravi Chellam, Kathleen
Conforti, Alain Dragesco-Joffe, Francisco Erize, Eskander
Ferouz, Helen Freeman, E.P. Gee, Mike Greer, Anne
Hilborn, Rafael Hoogesteijn, Lewis Horowitz, R. Idzerda,
Masako Izawa, Peter Jackson, David Jenny, Bholu Abrar
Khan, Sanjay Kumar, Claude Levinson, Paul Leyhausen,
Jill Mellen, Garth Mowatt, Kristin Nowell, Bharat Pathak,Dimitriy Pikunov, E. Ram&o, Tim Redford, D. Reucassel,
Kailash Sankhala, Roland Seitre, Gajandan Singh,
Alexander Sliwa, Shawn Smallwood, Fiona Sunquist,
Valmik Thapar, Jim Thorsell, Barbara Tonkin, Belinda
Wright and Giinter Ziesler. Thanks also to the Natural
History Museum, London, for the photo of a 19th century
painting of a Bornean bay cat by Joseph Wolf, and the
WWF Photo Library for photos by Tony Rath.

Masako Izawa (Japan); Hugo Jachmann (Malawi); Rodney
Jackson (snow leopard, management); Fabian Jaksic(Chile); Martin Jalkotzy (puma); A.J.T. Johnsingh (tigers,
India, research, management); Dereck Joubert (Botswana);
Dennis Jordan (Florida panther); Ullas Karanth (tigers,
India, research, management); Mohammed Khan (Malaysia); James Kirkwood (disease, reintroductions); Andrew
Kitchener (cat species, issues in cat conservation); Rick
Klein (Chile); Richard Kock (Kenya, disease); Gary
Koehler (Canada lynx, tiger research); Ashok Kumar(trade); Tor Kvam (Norway); Sally Lahm (Gabon); Karen
Laurenson (cheetah); Nigel Leader-Williams (Tanzania);
Paul Leyhausen (cat species, taxonomy); Donald Lindburg(cheetah in captivity); Malan Lindeque (Namibia); Fred
Lindzey (puma); Lu Houji (China); Ma Yiqing (China);
Kathy MacKinnon (Thailand); Stephen MacOrist (disease,
European wildcat); David Mallon (Ladakh and Mongolia);
Laurie Marker-Kraus and Daniel Kraus (cheetah); Rowan
Martin (leopard); E.N. Matjuschkin (former U.S.S.R.);
Charles McDougal (Nepal, management); Jeff McNeely(southeast Asia, issues in cat conservation);
Stephen
McOrist (disease); Heinrich Mendelssohn (Israel); Gus
Mills (South Africa, research); Fumi Mizutani (Kenya,management); Francis Mkanda (Malawi); Edgardo Mondolfi (Venezuela); Garth Mowat (Canada lynx); Elena
Mukhina (Central Asian republics); Iyad Nader (Saudi
Arabia); Stephen Nash (trade); Jan Nel (South Africa);
Howard Nelson (Trinidad and Tobago); John Newby(Saharan Africa); Peter Norton (leopard in South Africa);
John Oates (West and Central Africa); Stephen O’Brien(molecular genetics); U. Ohn (Myanmar); Gea Olbricht(cats in captivity); Madan Oli (snow leopard in Nepal,
management); William Oliver (Philippines); Craig Packer(lion); Bruno Paris (Guinea-Bissau); Junaidi Payne (Sabah
and Sarawak); Pierre Pfeffer (Bornean bay cat); Hubert
Planton (West and Central Africa); Kim Poole (Canada
lynx, trade); Howard Quigley (Latin America); Pat Quillen(cats in captivity); Alan Rabinowitz (tiger, clouded leopard, southeast Asia, research); Bernardino Ragni (Italy,
taxonomy); M.K. Ranjitsinh (India); Mohammed Reza
Khan (Bangladesh, United Arab Emirates); Wolfgang
Richter (Zaire); Tom Roberts (Pakistan); Robert Rolley(bobcat); Mark Rosenthal (flat-headed cat in captivity); Ian
Ross (puma); Jurgen Rottmann (Chile); Deb Roy (tiger,
India, management); Royal Forest Dept. of Thailand;
Richard Salter (Laos); Charles Santiapillai (Indonesia andVietnam); Pranabes Sanyal (India, management); Karen
Sausman (sand cat in captivity); Nan Schaffer (flat-headed
cat in captivity); George Schaller (Tibet); Lue Scheepers

XIntroduction

Introduction
summary sections which outline key points. A regional
index to species vulnerability, which generally indicatesspecies conservation priority, prefaces each regional chapter in Part I, the Species Accounts. The introduction to
the Species Accounts explains how species vulnerability
is ranked. Part III, the Action Plan, is organized according to the topics examined in Part II and the species orderof Part I.

The Cat Action Plan
Wild Cats: Status Survey and Conservation Action Planconsists of a review and analysis of information relevant tothe conservation of wild cats, and a priority action program. Part I provides summaries of the biology, ecology,
distribution, and conservation status of each cat species.
These Species Accounts are organized under five geopolitical regions: Sub-Saharan Africa, North Africa andsouthwest Asia, Tropical Asia, Eurasia, and the Americas.
Part II examines the major issues pertinent to the conservation of all cats: habitat loss, management of big cats nearpeople, research, trade, cats in captivity, and reintroduction. Parts I and II together form a comprehensive reference for people interested in cats and their conservation.
The information contained within is a demonstration of thework of cat specialists, and it is hoped that the rich andmulti-faceted picture of cats and their conservation which
emerges will stimulate more people to become active on
behalf of the wild cats.
Wild Cats is more, however, than an authoritative reference work. It is a strategic planning document whichprescribes methods for making cat conservation more
effective. These principles of cat conservation, which canbe drawn from the text, prioritize conservation action on
both international and regional levels. The principles also
serve as a framework to aid local authorities in planningtheir own cat conservation priorities.
Part III, the Action Plan itself, presents 105 projects thatbuild on the data and recommendations presented previously, and focus the general principles of cat conservation. Drawn up by the Cat Specialist Group, they
concentrate on the most vulnerable species and are priorities for cat conservation in the 1990s. Implementation ofthese projects forms the mission of the Cat Specialist
Group over the coming decade. If these projects realize
their objectives, the family Felidae should enter the 21st
century in good shape.
The priority projects listed in the Action Plan, for themost part, are in need of (1) financial support and (2)researchers and others to work on them. Those interested
in funding, carrying out, or helping with any of these projects should contact the Vice Chairman, Projects fordetails: Kristin Nowell, 2520-4,41st St. NW, Washington
DC 20007, U.S.A.
An Executive Summary of Wild Cats prefaces Part I. In
addition, the “Major Issues” chapters of Part II end in short

The Cat SpecialistGroup

The IUCNKSC Cat Specialist Group is the world’s premier body of scientific and practical expertise on wild cats
and their conservation. Over 160 members (see Appendix
5) represent 50 countries and include field biologists,
wildlife managers, government officials, leaders of nongovernmental organizations which focus on cat conservation, and other specialists from diverse but interrelated
fields including taxonomy, genetics, environmental law,wildlife trade and use, conservation education and wildlife
photography, small population biology and captive breeding, and wildlife veterinary medicine. These people serveas Cat Specialist Group members in their personal capacities, but bring with them the experience and the knowledge
gained in their professional careers. They volunteer thebest of their thinking, and also, in many cases, their timeand services, for cat conservation. This document represents the Group’s first major collective effort to review
what has been accomplished in the past, and to prepare a
strategic plan for future action.
Through its members, the Cat Specialist Group maintains a substantial collective library. The Group plans toconsolidate and disseminate this resource by establishing a
Cat Conservation Data Center (see priority project in Part
III). The Chairman publishes a biannual newsletter, Cat
News, which is circulated to members of the group. It isavailable to anyone else who makes an annual donation toa special fund in the name of “Friends of the Cat Group.”
For more information about the Cat Specialist Group,
contact: Peter Jackson, Chairman, IUCN/SSC
Cat
Specialist Group, Route des Macherettes, 1172 BougyVillars, Switzerland, Tel + Fax: +41 (21) 808 6012, email:peterjackson@gn.apc.org
or c/o the Species Survival
Commission, IUCN-The
World Conservation Union,
1196 Gland, Switzerland, Tel: +41 (22) 999 0001, Fax:
+41 (22) 999 0015, email: mgd@hq.iucn.ch (attn jackson).

xi

Executive

Summary

Executive

Summary
gal trade in tiger bone for Asian traditional medicines, andthis calls for expertise in several fields: protection of key
populations in reserves; protection of tigers outsidereserves; analysis of the consumer market for tiger bone;and effective enforcement of both national and international trade bans. This explains why more priority projects
are proposed for the tiger (14) than for the Iberian lynx
(3) in the Action Plan.
For many of the most vulnerable small cats, there islittle knowledge of their biology and ecology. Conservation of these species will be difficult without this baseline
information. There are thus relatively few projects forthese species beyond basic studies of natural history anddetailed surveys of the distribution of sub-populations.

Wild Cats: Status Survey andConservation
Action Plan
There are 36 species of wild cat, ranging in size from thetiger to the tiny rusty-spotted cat. They are found in every
continent except Australia and Antarctica. This document, Wild Cats: Status Survey and Conservation Action
Plan, is designed to promote the conservation of all thewild cats in their natural surroundings. The increase in
numbers of people, the spread of settlement and the
exploitation of natural resources of wild lands hitherto
little disturbed, together with persecution, are threateningsome cats with extinction. Other cat species are declining in numbers.
To assess the vulnerability of the cats, a system has
been developed to rank them in five main categories, bothon a world basis and a regional basis. The ranking system is based on four factors:

Part I: Species Accounts
The Species Accounts provide the latest information on the
biology, ecology and conservation status of the wild cats.
Photographs illustrate the characteristics of each cat, andmaps provide the latest information on their range. The
species accounts serve as a database, to be built on as
research produces more information.

1. The number of habitat types with which each species
is associated: the fewer habitats with which a species
is associated, the more vulnerable it is to habitat loss.
2. Total range size: the smaller the distribution of a species,
the more vulnerable it is to further loss of range.
3. Body size, which provides a link to esti mates of total
numbers: the larger the cat, the fewer the number ofindividuals likely to be located in a given area compared with smaller cats.
4. Active Threat (“A”), which refers to high levels of
hunting pressure, leading to the loss of animals fromhabitat in which they would otherwise be present.

Part II: Major Issues in Cat Conservation
Part II is devoted to the review and analysis of six major
issues in cat conservation:
Chapter 1.
Chapter 2.
Chapter 3.
Chapter 4.
Chapter 5.
Chapter 6.

Combining the scoring on these factors makes it possible
to group cats into categories in order of their priority forconservation (Boxes 1 and 2). The ranking system is
described in detail in the introduction to Part I.
In general, species in Category 1 warrant first attention, both globally and regionally. Yet practical considerations play a large role in determining how conservation
action should be structured in terms of projects. For example, the Iberian lynx emerges as the most vulnerable wild
cat and is ranked ahead of the tiger, although the tiger isseriously threatened. However, it will take much moreconservation effort to save the tiger than the Iberian lynx.
The total numbers of Iberian lynx may be fewer than tiger,
but the lynx occurs mainly in Spain, and this allows forunified conservation action. The tiger is scattered in small,
localized populations in 14 Asian countries, including theworld’s two most densely populated nations, India andChina. Moreover, the primary threat facing the tiger is ille-

Habitat loss and fragmentation
Management of big cats near people
Research
Trade
Captive breeding
Reintroduction

Examination of these issues leads to several inter-related
conclusions about cat conservation, which are summarizedat the end of each chapter. These are reviewed in thisExecutive Summary in the form of key general questions
which the Action Plan projects are designed to answer.
How do cat species adapt to dlj$erentforms ofhabitat loss and modification?
Chapter 1 shows that most of the world’s original naturalvegetation has been modified in some way by people.
Protected areas cover only small portions of the range ofmost species, so that most cats live in modified habitat.
Fortunately, cats, not having specific vegetation requirements, are more flexible than many other animals in termsxiiExecutiveof their ability to persist. For example, logging in tropicalrain forest doesnot necessarilyleadto the declineor extirpation of its cat populations. As discussedin Chapter 3,
there have been relatively few studies of cats in alteredhabitats;mosthave beendone in protected areas. Several
Action Plan projects (numbers 2 and 23) have been
designedto identify the variableswhich permit catsto persist in different forms of modified habitat. In addition, anumber of natural history studies are proposed whichshouldbe carried out both in good-quality protected habitat and in a type of modified habitat which predominates
within the range of the species. It is important for thesestudiesto estimatecat densitiesin modified habitat, so as
to calculatenumbersover large areasof their ranges.

Box 1
Global Ranking of Cat Species
Vulnerability
Category 1 (Top priority)
lberian lynx

lynx pardnus

Category 2
Tiger (A)Snow leopard (A)Bomean bay cat
Chinese mountain cat
Black-footed cat
KodkodAndean mountain cat
Flat-headed cat
Fishing cat
African golden cat

Panthera tigris
Uncia uncia
Catopuma badia
Felis bieti
Fe/is nigripes
Oncifelis guigna
Oreailurus jacobitus
Priunaiiurus planieeps
Prionailurus viverrinus
Profelis aura ta

Category 3
Cheetah (A)Lion (A)Jaguar (A)Asiatic golden cat
OncillaRusty-spotted cat
Clouded leopard
Marbled cat

Acinonyx jubatus
Panthera lea
Panthera onca
Catopuma temmincki
Leopardus tignnus
Prionailurus rubiginosus
Neofelis nebulosa
Pardofelis marmorata

Category 4
Sand-catMargayServalCanada lynx
Geoff roy’s cat
Manul

Fe/is margarita
Leopardus wiedi
1eptailurus serval
Lynx canadensis
Oncifelis geoffroyi
Otocolobus manul

Category 5a
Puma (A)
Leopard (A)OcelotEurasian lynx
BobcatPampas cat

Puma concolor
Panthera pardus
Leopardus pardalis
lynx lynx
Lynx rufus
Oncifelis colocolo

Category 5b
CaracalJungle cat
Leopard cat

Caracal caracal
Fe/is chaus
Prionailurus bengalensis

Category 5c
WildcatJaguarundi

Fe/is s/Ives tris
Herpailurus yaguarondi

Summary

Which types of habitat are mostimportantfor cat speciesconservation?
Certain habitat types are the richest in vulnerable cat
species,but are either declining in areaor becomingfragmented. Theseinclude tropical moist forest, especiallyin
the lowlands, tropical montane complexes, high alpine
habitat, andmajor wetlands. Conservationof thesehabitat
types is important for cat species.Protectedareasneedto
be sufficiently large to support viable populations of thebiggestcats.
What managementmeasurescan be taken to promoteconservationof big cats living near people?
Most catsare found outsideprotected areas,and live nearpeople. They risk extirpation through unsuitablemodification of their habitat, depletionof their prey, and persecution. This is particularly true for big cats,which can causesignificant economiclosseswhen they prey on livestock,particularly for poor owners of a few animals. Several
Action Plan projects focus on the study of big cats whichlive amongpeople,andon the recommendationandimplementationof effective managementmeasures(Projects56,
22,23, 31, 32,40,5 1,52,71,74,75,93, and 102).
What are the biological and ecologicalrequirementsof vulnerable cat species?
To evaluate the conservation status of cat specieson anational or regional scale, a basic understandingof their
biology and ecology is needed. Otherwise, it is difficult
to plan specific conservationmeasures.For a surprisingly
high number of vulnerable species,natural history has
never been studied,either in detail or at all. A numberof
Action Plan projects have been put forward to addressthese major gaps in our knowledge of the cat family(Projects 2, 18, 20, 34, 37,42,43, 57, 58, 60, 61, 63, 64,
66, 68, 76, 77, 91, 92, 94, 96, and 99). In addition, longterm studieswhich have gatheredcomprehensivebaseline
dataon cat populationsshouldbe continued.

A=Actively Threatened(High levels of hunting pressure)
See pages 2-6 for exptanation ofvulnerability ranking system.

m..
XIII

Executive

Summary

Box 2
Regional Ranking of Cat Species Vulnerability
Sub-Saharan Africa
1. Black-footed cat
1. African golden cat
2. Cheetah (A)2. Lion (A)3. Serval
4. Leopard (A)
4. Caracal
5. African wildcat

North Africa and Southwest
1. Cheetah (A)1. Asiatic lion
2. Serval (A)3. Leopard (A)4. Sand cat
5a. Caracal (A)5a. Jungle cat
5b. Wildcat

Tropical Asia
1. Tiger (A)I, Bornean bay cat
2. Clouded leopard (A)2. Asiatic golden cat
2. Flat-headed cat
2. Rusty-spotted cat
2. Fishing cat
2. Marbled cat
3. Leopard (A)4. Jungle cat
5. Leopard cat

Eurasia
Asia sub-region
I, Snow leopard (A)1. Chinese mountain cat
2. Asiatic wildcat
2. Manul
3. Eurasian lynx

Asia

The Americas
1. Kodkod
1. Andean mountain cat
2. Jaguar (A)
2. Oncilla
3. Margay
3. Canada lynx
3. Geoffroy’s cat
4. Puma (A)
4. Bobcat
4. Pampas cat
5. Jaguarundi

Europe sub-region
1. lberian lynx
2. Eurasian lynx
3. Eurasian wildcat

Note: lriomote cat not ranked but high priority (see page xiv).
See pages 2-6 for explanation of vutnerabitity ranking system.

tory of tiger censusesand is home to most of the world’s
tigers. The improved techniqueswill be applicablein thecensusof other big cats.

How fragmented are cat speciespopulations?
Many cat specieshave been extirpated from large partsof their ranges, but this has rarely been mapped.
Population fragmentation can result in small, isolated
populations, which are particularly vulnerable to extinction. Mapping the detailed distribution of cat populations
will greatly aid in fixing priorities for the conservation
effort, anda numberof Action Plan projectshave beenput
forward to start this process(Projects 3,4, 19,21, 2526,
28, 39, 41,42, 49, 53, 59, 62, 65, 72, 73, 80, 90, 91, 92,
97,99, and 104).

How can the viability ofcatpopulations be ensured?
Studiesin conservation biology show that small, isolated
populations are highly vulnerable to decline and extinction. Becausecats occur at relatively low densities,mostprotected areas conserve only small populations. Are
thesepopulations too small to be viable? Project 8 aims
to apply the MVP concept to cat species,particularly thelarger cats, in the light of what is known of their biology
andecology. This information will thenbe usedto analyze
the viability of sub-populationsthroughout the range of a
species.Other aspectsof MVP conservationare covered,including the extent to which habitatcorridorscan enhance
the persistenceof a populationby allowing the movementof individualsbetweenpopulations(Project 50); the role ofdiseasein smallpopulations(Projects 11 and 27); limiting
factors of populations (Projects 36,46, and 55); and theecology of isolatedpopulations,particularly in termsof the
impact of predation on numbersof prey (Project 29).

How can cats be counted effectively?
Becausecats are mainly nocturnal and secretive, they are
difficult to census. But it is impossibleto assessthe status of a speciesin a given areawithout reliable indexesof
numbersandpopulationtrends. This is particularly important for the tiger, which is beingheavily poachedfor bones
andother parts. Two Action Plan projects(numbers7 and50) are designedto develop and promote appropriatecensusmeasures. One of them focuses on improving thecounting technique used in India, which has a long his-xiv

Executive

How can the conservation of’11 intraspecificdiversity be ensured?
There is general agreement that most classically describedcat subspecies are not valid, but little progress has been
made in re-defining subspecies using modern techniques,
including genetic analysis. The Action Plan identifies taxa
for which both field conservation efforts, as well as genetic
studies, are of highest priority (Projects 10, 36,44,46,47,
56, 67, 69, 70, 78, 79, 89, 95, 100, and 105). In addition,
it is recommended that field biologists increase theirefforts to collect biological samples to help in evaluating
subspeciation (Project 9), and that zoos continue theirefforts to identify subspecies and conserve viable populations of key taxa (Project 15).
How can illegal trade in cat products becontrolled efectively?
Commercial poaching and illegal trade are serious threats
to some species. To minimize them, a great deal of information is required about the consumer market for cat products. There is a need to know about the size of the market,
sales volume, trade channels and patterns, key players in
the market, consumer motivation, and law enforcement
measures and their effectiveness. This is most urgent for
the tiger: the consumer market for tiger bone medicines
has scarcely been investigated (Project 12).
How can the sustainability of hunting
for cats be ensured?
Economic value is an important incentive for cat conservation, and some of the most significant values are derived
from hunting for the fur trade and for sport. Projects areproposed to further develop management techniques toensure that hunting pressure does not lead to majordeclines in numbers and that yields are sustainable(Projects 13, 33, and 98).
How can zoos contribute most efictively
to cat conservation?
In terms of captive cat populations, experience in thegenetic management of small populations, and the ability

Summary

to promote cat conservation, zoos have resources which
are becoming increasingly appreciated by the conservation
community. Expertise in small population biology is called
for in Project 8. In addition, as zoos become involved withconservation of wild populations, a zoo-sponsored fund
for field conservation is proposed (Project 14).
How well does cat reintroduction work, andfor
which taxa is it a priority?
Reintroduction of captive-bred cats is often seen as ameans of maintaining wild populations. However, reintroduction is a complicated matter and is not practical if
the factors which led to the decline or extinction in the firstplace have not been alleviated. The establishment of a
population can be difficult when habitat is fragmented andused by humans. Projects 16, 17, 84, and 85 monitor the
long-term progress of reintroduction attempts. In general,
reintroduction is not of high priority for cats, because none
have become extinct in the wild, and efforts should firstbe focused on ensuring that they do not. However, theAsiatic lion is in a grave situation because it survives only
as a single, highly vulnerable wild population in India’s
Gir Forest. As insurance against sudden, catastrophic
extinction, a second population urgently needs to be established. Possible sites are being surveyed and assessed inIndia (Project 35).

Part III: Action Plan
Members of the Cat Specialist Group are involved in studies and conservation of cats in all the continents. They
have provided information about their current projects,
and proposed others that they consider priorities for conservation in the 1990s. Other projects have arisen fromthe research conducted to produce this document. Part
III provides summaries of 105 projects. Some alreadyhave financial support, but most require funds if they are
to be implemented. The Cat Specialist Group will actively
seek funding for these priority projects, and hopes for
sympathetic consideration by major institutions, as well asprivate donors.

Taxonomyof the Felidae

Taxonomyof the Felidae

In practice, conservation of cats in the wild has to be basedon populations rather than taxonomy, but taxonomy is an
aid to prioritizing allocation of conservation resources
between different populations. It is thus essential that classification schemes accurately capture felid diversity, interms of not only morphology, but also genetics, and, if
possible, ethology. The history of cat species classification, which has seen extremes in both “splitting”
and“lumping,” is reviewed below by Lars Werdelin in a paper
written especially for this volume.
Wild Cnts follows the taxonomy set out in the latest edition of Mammal Species of the World (Wozencraft 1993).
Wozencraft has explained that his taxonomy is not a pieceof primary research, but a compilation of recent literature.
He evaluated what others had done, based on primary literature, discarding statements unsupported by data (C.
Wozencraft in Zitt. 1993). His classification is used here
for practical reasons, without prejudice, as it has been
adopted by the Convention on International Trade inEndangered Species of Wild Fauna and Flora (CITES) andthe World Conservation Monitoring Centre (WCMC).
Some of the new designations are controversial and will
surely be the subject of future debate. One example is the
Iriomote cat, which was originally described as a monotypic genus Mayailurus iriomotensis (Imaizumi 1967), but
was later placed close to the leopard cat within the genus
Prionailurus (Hemmer 1978a, Leyhausen 1979, Corbettand Hill 1993): it is now relegated to a subspecies of theleopard cat by Wozencraft (1993). If the Iriomote cat is
considered a full species, it is the most endangered cat in
the world, with a population of only 100 animals on thesmall Japanese island of Iriomote. If it is considered a subspecies, it becomes one of several island populations of themost common cat in Tropical Asia. Nevertheless, becauseof its distinctive characteristics, which led to the uncertainty in classification, the Iriomote cat merits special
attention.
Modern taxonomic frameworks have lumped most of
the smaller cat species into the genus FeZisaccording to the previous edition of Mammal Species qf theWorld (Nowak and Paradiso 1983). Wozencraft (1993),
however, broadly promotes the subgenera of the old genus
FeZis to full generic status, a step which better reflects thesubstantial variation among so many species. As Pocock( 195 1) noted in his Catalogue of the Genus Felix “[Theold genus] Felis is a heterogeneous, unwieldy assemblage,
ranging practically all over the world, apart from
Madagascar, some small islands and the Australian
Region. Considering its wide distribution and exceedingly
varied habitats, it would be strange if the family had not

become differentiated into groups of generic status.”
Under the new taxonomy, a number of former Felis
species are now placed in monotypic genera: the caracal,
jaguarundi, serval, Andean mountain cat, Pallas’s cat,
African golden cat, puma, and marbled cat. The threelynxes-Eurasian(lynx), Canada (canadensis), and Iberian(pardinus)-oftenlabelled conspecific, have full species
status within the genus Lynx. The strongly-patterned spotted South American cats-ocelot (pardalis), oncilla (tigrinus), and margay (wiedi)have
been placed in the genus
Leopardus, while the lightly spotted cats of the southern
region-pampas cat (coEocoZo), Geoffroy’s cat (geoflroyi),
and the kodkod (guigna)-aregrouped in the genus
Onc$elis. The Asiatic golden cat (temmincki) and the
Bornean bay cat (badia) are placed together in the genus
Catopuma. The snow leopard is separated from the genus
Panthera and given full generic status as Uncia uncia.
With regard to subspecies, there is considerable debate
on definition, and even whether the traditional taxonomic
concept is valid in the light of contemporary knowledgeof population biology and genetics. It is generally agreed
that too many subspecies of cats have been described in thepast on the basis of very slim evidence. However, there issome uncertainty as to how to proceed with redefinition,and the task is large-the Felid Taxon Action Group of theAmerican Zoo and Aquarium Association (formerly the
AAZPA), which is concerned mainly with appropriate representation of wild diversity through captive breeding, hasrecommended the re-evaluation of 235 out of 259 subspecies recognized by the group (Wildt et al. 1992a).
Molecular analysis is potentially an important tool forthis exercise. The leopard, for example, shows great variation in coat pattern and size, but recent molecular studies
have led to the proposal that all African leopards-nearly
30 have been described, living in habitats which range
from desert to rain forest-should
be considered one subspecies Panthera pardus pardus (Miththapala 1992).
However, can such findings be reconciled with data gathered by classical anatomical measurements and descriptions, and with what new knowledge has been gained
through field studies of behavior and ecology in different
environments? In Part II Chapter 3, where the questionof subspecies is discussed in more detail, Stephen O’Brien
puts forward a useful definition, outlines the sort of evidence of differentiation that molecular biologists shouldlook for, and stresses the need for cooperative work
between the different scientific disclipines.
Given the difficulty of defining subspecies and the lackof comprehensive evaluation at this level of the family
Felidae, this Action Plan refers only to those subspecies

xviTaxonomy

Box 1Classificationof the Felidae

by W. Christopher Wozencraff (1993)
Family Felidae G. Fischer, 1817
Subfamily Acinonychinae

Pocock, 1917

Acinonyx Brookes, 1828
jubatus Schreber, 1776

Cheetah

Subfamily Felinae Fischer, 1817
Caracalcaracal (Schreber, 1776)

CaracalCatopuma Severtzov, 1858
badia (Gray, 1874)
temmincki 1 (Vigors and Horsfield, 1827)

Bornean bay cat
Asiatic golden cat

Fe/isbieti Miine-Edwards, 1892
chaus Schreber, 1777
margarita Loche, 1858
nigripes Burchell, 1824
silvestris Schreber, 1775

Chinese mountain (desert) cat
Jungle cat
Sand catBlack-footed cat
Wildcat of Africa and Eurasia

Herpailurusyaguarondi Lacdpbde, 1809

Jaguarundi

Leopardus Gray, 1842
pardaiis (Linnaeus, 1758)
tigrinus (Schreber, 1775)
wiedi (Schinz, 1821)

OcelotOncilta, Little tiger cat
MargayLeptailurus Severtzov, 1858
serval (Schreber, 1776)

ServalLynx Kerr, 1792
canadensis Kerr, 1792
lynx (Linnaeus, 1758)
pardinus (Temminck, 1824)
rufus (Schreber, 1776)

Canada lynx
Eurasian lynx
Iberian lynx
BobcatOncifelis Severkov, 1858
colocoio (Molina, 1782)
geo#Voyi (d’urbigny and Gervais, 1844)
guigna (Molina, 1782)

Pampas cat
Geoff roy’s cat
KodkodOreailurus Cabrera, 1940
jacobitus (Cornalia, 1865)

Andean mountain cat

Otocolobus Brandt, 1842
manul (Pallas, 1776)

PaHas’s cat

Prionaiiurus Severtzov, 1858
bengalensis (Kerr, I 792)
planiceps (Vigors and Worsfield, 1827)
rubiginosus (1. Geoffroy Saint-liilaire, 1831)
viverrinus (Bennett, 1833)

Leopard cat
Flat-headed cat
Rusty-spotted cat
Fishing cat

xvii

of the Felidae

Taxonomyof the Felidae

Profelis Severtzov 1858.
aurata (Temminck,

1827)

African golden cat

1771)

Puma, Cougar, or Mountain

PumaJardine, 1834.
concolor (Linnaeus
Subfamily

Pantherinae

Pocock1917

Neofelis Gray, 1867
nebulosa (Griffith, 1821)

CloudedPanthera Oken, 1816.
leo (Linnaeus, 1758)
onca (Linnaeus, 1758)
pardus (Linnaeus, 1758)
tigris (Linnaeus, 1758)

Lion
JaguarLeopardTiger

Pardofelis Severtzov, 1858
marmorata Martin, 1837

MarbledUncia Gray 1854
uncia (Schreber,

Snow leopard

1758)

lion

leopardcat

Wozencraft, W.C. 1993. Order Carnivora. Pp. 286-346 in D.E. Wilson and D.M. Reeder, eds. Mamma/speciesof the world: a taxonomic and geographic reference (Second edition). Smithsonian Institution Press, Washington
D.C. and London.
1 jacobita, wiedii, and temminckii in Wozencraft (1993) amended to jacobitus, wiedi, and temmincki in accordancewith the 1985 International Code of Zoological Nomenclature Article 31a mandating that patronymic species names
follow the rules of Latin grammar.
Note: Brackets round the name of the authority indicate that the genus has been changed
publication by that authority.

which have beenrelatively rigorously evaluated,and generally prioritizes conservation action at the specieslevel.
However, it is also recognized that preservation of a
speciesincludes its full diversity, and that at presentit isintra-, rather than interspecific diversity, which is mostthreatenedin the cat family. A list of classicallydescribedsubspeciesis included in Appendix 1, and much greater
effort shouldbe directed towards defining differentiation
within cat speciesthrough more extensive collection andanalysisof field samples.A protocol for the collection offield samples,a collaborative effort by a museum-based
anatomist and a wildlife veterinarian, is contained inAppendix 2. There area numberof taxonomy-related priority projectsin Part III.
Below, Lars Werdelin reviews historical efforts toclassify cat species,and Stephen O’Brien discussesthe
usefulnessof genetic analysis in clarifying felid evolutionary history.

...
XVIII

The History

since firstof Felid Classification

by Lars Werdelin
Like most other groups of organisms, felids have been
the subject of a number of revised classifications since
Linnaeus (1758), in the 10th edition of his Systema
Naturae, laid down the first foundations by naming the
genus Felix The following is an attempt to provide abrief history of theseclassification attempts, illustrating
our growing understanding of the interrelationships ofthe living cats. In addition to the works discussedhere,
which are all mainly concerned with living felids, there
have been many works that in a general way have tried
to relate fossil and living felid taxa. However, most ofthesehave not dealt specifically with the ancestorsof living species,or where they have done so, have concerned
themselves with only a limited set of taxa. Werdelin(198 1) is an example of such a study. These have notTaxonomyof the Felidae

lynx, Canada lynx, and bobcat, and Urolynchus
for thecaracal.
5. Felis, which contains no less than 19 subgenera, mostly
monotypic.
Oncoides: ocelot, margay, and oncilla;
Pardofilis: marbled cat;Catopuma: Temminck’ s golden cat;
Herpailurus: jaguarundi;
Leptailurus: serval;
Chrysailurus: for one variety of theAfrican golden cat;
CatoZynx: domestic cat (in which he presumably
included the European wildcat), the African
wildcat, and the jungle cat;
Otocolobus: manul;
Lynchailurus: pampas cat;
Oncifelis: Geoffroy’s cat;
Noctifelis: kodkod;
Profelis: another variety of the African golden cat
Dendrailurus, which is based on an unidentifiable
species;
Fe&, which is preoccupied by Linnaeus’ FeIis forthe domestic cat;Prionailurus: leopard cat;
Zibethailurus: fishing cat;
Ictailurus: flat-headed cat;
Otailurus, for a species from Timor that I am
currently unable to identify.

been considered in this review.
The first author specifically to consider relationships
between species within the family Felidae was Jardine(1834).
He distinguished five genera, Leo, Puma,
Cynailurus, Lynchus, and Felix In the first of these heplaced only the lion, then separated into two species. In
the second he had the puma, the jaguarundi, and the pampas cat (one color phase). In Cynailurus he placed only the
cheetah. In Lynchus he placed the Eurasian and Canada
lynxes, as well as the bobcat, caracal, African golden cat,
Geoffroy’s cat, jungle cat and black-footed cat. In Felis,
finally, he placed all other species known at that time:
tiger, leopard, jaguar, snow leopard, ocelot, margay,
oncilla, leopard cat, clouded leopard, serval, pampas cat(other color phase), and European, African, and Asian wildcats. He did not consider relationships within these genera.
Although quite different from our current conception of
felid interrelationships, Jardine’s classification nevertheless contains some themes which have run through thesubject ever since. These include the recognition of agenus Felis sensu strict0 (although broader than currently
conceived); the recognition of Lynx as a distinct genus(also broader than currently conceived); the relationshipbetween the caracal and the lynxes; and the relationshipbetween the puma and the jaguarundi. The latter relationship, which is highly controversial, has been supported bymany authors since, and it is interesting to find its roots atsuch an early stage of the game.
Jardine was a precursor and his classification a crude
first attempt, although an interesting one. The modern
age of felid classification begins with Severtzov (18571858). This author discussed the evolution of carnivores ingeneral and felids in particular, with special emphasis on
biogeography and its relationship to felid classification. In
the process of so doing he erected a number of new genuslevel names as subgenera. In total, his classification
includes five genera and 27 subgenera. Severtzov’s exposition is not easy to follow, perhaps because he hadplanned to follow this work by a more extensive monograph on the group, where he intended to publish the characteristics of his various groups. This work was apparently
never published. Fortunately, Allen ( 19 19b) published aclarification of Severtzov’s concepts, considerably simplifying a review.

This enormous proliferation of generic-level names
clearly does nothing to increase our knowledge of theinterrelationshipsof the various species. However, it
should be noted that most of the names used by Severtzov,whether newly coined by him or adopted from earlier
authors, are still in use for various groupings of felid taxa.
In Severtzov’s classification we see the seeds of a modern
concept of Panthera in his genera Panthera and Tigris.
His concept of Lynx is also very close to the current one.
His Oncoides represents the beginnings of the currently
recognized Leopardus for the small spotted felines ofSouth America. Other than this, Severtzov’s contribution is mainly at the nomenclatural level, albeit a very
modern one.
While Severtzov was publishing his work, Gray (1867)
was completing his studies of felid classification. Gray
was apparently unaware of Severtzov’s work, and therefore there is extensive overlap between them, as well as anumber of synonymous taxon names. In Gray’s classification, the pantherines are separated into four genera:
Uncia for the snow leopard; Leo for the lion; Tigris forthe tiger; and Leopardus for the leopard, jaguar, African
golden cat, and puma. This is one of the few notions thatthe golden cats are related to the pantherine big cats. The
genus Neofelis includes the clouded leopard, whereas the

Severtzov’s genera are as follows:
1. Tigris, which includes two subgenera, Leo for the lion
and Tigris for the tiger.
2. Panthem, with the subgenera Jaguarius for the jaguar,
Panthera for the leopard, Uncia for the snow leopard
and clouded leopard, and Puma for the puma.
3. Cynailurus, with the single species Cynailurus jubatus, the cheetah.
4. Lynchus, with two subgenera: Lynchus for the Eurasianxix

Taxonomyof the Felidae

genus Pardalina includes an unidentified species, P.
himalayensis, possibly an ocelot. The genus Catolynx inGray’s conception includes only the marbled cat. This
genus is therefore synonymouswith Severtzov’s
Pardofelis, but is itself a junior homonym of CatoZynx ofSevertzov, which is a junior synonym of Felis sensu strict0(they are based on the same type species). This chain reaction is a good illustration of why the parallel work of
Severtzov and Gray has led to over 100 years of nomenclatural confusion in felids. No wonder many workers
take refuge in calling everything Felis.
Gray’s genus Viverriceps includes the fishing cat, flatheaded cat, rusty-spotted cat, and one variety of leopard
cat; his genus Pajeros includes only the pampas cat. In thegenus Felis Gray places the ocelot, margay, oncilla,
Geoffroy’s cat, jaguarundi, serval, Asiatic golden cat,manul and the European, African, and Asian wildcats,
along with the domestic cat. In his genus Chaus he places
the jungle cat, while in Lyncus he has the Eurasian,Canadian, and Iberian lynxes, and the bobcat. His genus
Caracal accounts for the caracal, while in Gueparda,
finally, he places the cheetah.
As noted, Gray’s work introduced some confusion in
the nomenclature, but he is more specific regarding interrelationships than Severtzov, for whom most species
belonged in their own genera. Gray’s genus Viverriceps,
for example, is a specific statement of relationships
between four species of southeast Asian felid. His conceptof Lynx is the same as that currently in use. On the other
hand, his Felis includes both species currently placed inthat genus and a number of species currently believed to be
only distantly related to Felis sensu stricto.
Some semblance of order was created out of thenomenclatural confusion by Pocock (19 17), who has perhaps done more than any other biologist to further thecause of felid classification and systematics. He separated
the Felidae into three subfamilies: Felinae for the smallcats, Pantherinae for the large (roaring) cats, andAcinonychinae for the cheetah. This classification was
based on the structure of the hyoid (ossified in Felinae andAcinonychinae, imperfectly ossified in Pantherinae) andthe digits (cutaneous lobe protecting retracted claw inFelinae and Pantherinae, no cutaneous lobe in Acinonychinae). This is a scheme which, with few exceptions,has been followed until very recently.
Within the Pantherinae, Pocock distinguished two genera: Panthera for the lion, tiger, leopard, and jaguar; andUncia for the snow leopard. Within the Felinae, he tried toarrange Severtzov’s and Gray’s genera in an orderly manner. Pocock’s Felis includes, in his terms, “three categories”: medium-sized cats from Europe, southwest Asia,
and Africa (these are not specified, but presumably include
the European, Asian, and African wildcats); larger species
ranging from Burma, through India, and into parts of cen-tral Asia (this group he specifically states is identical with
Gray’s genus Chaus, i.e., the jungle cat); and the verysmall South African species F. nigripes, the black-footed
cat. As Pocock is no more explicit about the species ofFelis than this, it is not clear where he placed the sand cat,
Chinese mountain cat, etc. relative to these three groups.
Within Lynx Pocock also distinguishes three groups:
one for the Eurasian, Canada, and Iberian lynxes; one forthe bobcat; and one for the caracal. He places the manul inthe genus Trichaelurus, the puma in the genus Puma, andthe serval in the genus Leptailurus. In Prionailurus heincludes both the leopard cat and the rusty-spotted cat,while in Pardofelis he places the marbled cat and the
Bornean bay cat. The genus Profelis includes both the
African and Asiatic golden cat, the first association ofthese two species.
His genus Zibethailurus includes the fishing cat, while
ktailurus
includes the flat-headed cat. Neofelis includes
the clouded leopard, while Leopardus includes only theocelot and margay. The other small South American cats(excepting the pampas cat, which Pocock identifies withthat species made the type species of Dendrailurus by
Severtzov) are included by Pocock in Herpailurus, whichaccordingly accommodates the jaguarundi, kodkod,
Geoffroy’s cat, and oncilla.
In summary, Pocock’s genera are to a great extent congruent with those recognized at present. His Panthera,
Felis, and Lynx (almost) are those currently in use, as aremany of his smaller groups. However, Pocock’s aim wasstrictly a classification, and he did not go beyond thisscheme to look at the interrelationships of the groups heproduced. This somehow led to the impression that there
were no such interrelationships to be obtained from thedata, and this, coupled with the massive influence ofPocock’s work, caused research on felid classification andsystematics to grind to a halt for more than half a century.
During this hiatus there were no studies emphasizing
felid classification. Some works, such as that of Weigel(196 1). include evolutionary schemes for the Felidae thatcan be made into classifications, but this was not theirmain aim. Finally, Hemmer (1978a) produced a considered view of felid interrelationships.
Hemmer also provides a phylogenetic tree, which none of the older workers
did. Therefore, his scheme of relationships, and by extension his classification, is more explicit than those of
Severtzov, Gray and Pocock (Fig. 1).
Hemmer considers the genus Felis sensu strict0 to bemonophyletic, incorporating the European, African, andAsian wildcats, which are considered closely related, andthe black-footed cat, Chinese mountain cat, sand cat, andjungle cat. Related to these are also the manul, placed in
the genus Otocolobus, and the lynxes, Lynx, the caracal,
Caracal, and the serval, Leptailurus.
The genus
Prionailurus is extended in Hemmer’s scheme to include

Taxonomy

Figure1. Branching

diagramderivedfrom the phylogenetic

tree proposed

by Hemmer(1978).

Source:Sallesof the Felidae(1992).

cats (Leopardus), the Felis sensu strict0 lineage (including
the manul), and the pantherine lineage with Lynx as the sister group of Panthera.
At the same time as Collier and O’Brien, Herrington(1986) prepared a systematic study and classification of
felids (Fig. 2), with partially congruent results. She also
has Panthera and Lynx closely related, although separated
by the marbled cat and clouded leopard. Herrington further recognizes Leopardus in more or less the same way as
Collier and O’Brien, although she considers Profelis,
including the golden cats and the Bomean bay cat, closely
related to the South American group. Herrington also recognizes Felis sensu stricto, but has the caracal and rustyspotted cat as close relatives of this genus. She sees thecheetah, jaguarundi, manul, and puma as closely related,
and identifies a genus Prionailurus including the fishingcat, leopard cat, flat-headed cat, and Iriomote cat.
It is noteworthy that the three assessments by Hemmer,
Collier and O’Brien, and Herrington all depart more or lessstrongly from the threefold subfamilial division-thebigcats, the small cats, and the cheetah-espoused by Pocock.
The most recent studies depart strongly from this scheme

the Iriomote cat, flat-headed cat, leopard cat, rusty-spotted cat, and fishing cat, of which the last three are considered more closely related. The small South American
forms are separated-into Leopardus for the ocelot and margay, Lynchailurus for the pampas cat, Oreailurus for the
Andean mountain cat, and Oncifelis for the oncilla,
Geoffroy’s cat, and kodkod.
Hemmer’s Panthera includes the same four species as
Pocock’s, but he considers the tiger more distantly related
than the other three, along with the snow leopard, Uncia. In
the large cat clade he also has the clouded leopard, NeofiZis,the marbled cat, Pardofelis, andxthe African golden cat,
Profilis. In Catopuma Hemmer united the Bomean bay cat
and Asiatic golden cat. The cheetah is alone in Acinonyx,
while he sees a close relationship between the puma, in
Puma, and the jaguarundi, in Herpailurus.
The next step in felid systematics and classification wasessentially twofold. In 1985, Collier and O’Brien published the first molecular systematic study of the Felidae,
with a number of innovative results (see next paper). My
view of classification essentially follows theirs, with athreefold division into the small South American spottedxxi

Taxonomy

Figureof the Felidae

2. Branching

diagramredrawnfrom the cladogramfor extant

felids proposedby placing the cheetah well within the felid radiation,
instead of as a separate lineage as was done by the earlier
workers, including Hemmer.
Another recent work in the field of felid systematics
and classification is that of Salles (1992). His study recognizes two well-resolved
groups and a basal group ofless well understood taxa (Fig. 3). One of the wellresolved groups includes the genus Felis sense stricto,
which in Salles’ view includes the manul. This genus isrelated to Lynx-, including the caracal. In this larger group
we also have the marbled cat, Bornean bay cat, andAsiatic golden cat.
Salles’ second large group is the pantherine group,
which beside Panthera also includes the clouded leopard,
cheetah, snow leopard, puma, and jaguarundi. The rest ofSalles’ relationships are basically unresolved, but we may
note the suggestion that the flat-headed cat and fishing cat
are closely related and are the basal felid group.
Wozencraft’ s ( 1993) classification, used in this volume,is the most recent evaluation of the felid family. He recognizes three suprageneric groups: the Acinonychinae
for the cheetah, the Felinae for the small cats, and theby Herrington(1986).

Source:Sales(1992).

Pantherinae for the large cats. He includes Neofelis andPardofelis in the Pantherinae along with Panthera andUncia, which reflects the opinion of some workers (i.e.,
Hemmer) that the small marbled cat is actually closely
related to the large cat group.
This survey represents a sample of the work on felidclassification and systematics undertaken during the past
>150 years. It is by no means complete, not taking into
account work by authors such as Matschie, Satunin,
Groves, Kratochvil, and others. However, a complete
review would occupy far too much space, and this brief
overview is more or less representative of the diversity ofviews on the subject.
What can we learn from this history? I feel that there
are two things that need to be pointed out. The first is thatthe divergence of opinion regarding felid systematics
expressed in even the most recent works suggests that considerable further work is required before a stable consensus can be reached. Such a consensus must involve bothmorphological and molecular work. The second important
point to be learned is that nearly all first-hand studies of
felid systematics and classification have separated felidsxxiiTaxonomyof conservation leads not only to confusion in establishing management plans, but risks critical mistakes in establishing priorities in cases where taxonomy is based oninadequate descriptions with only historic precedence toaffirm their precision. In the past decade conservation
efforts for several species have been both advanced andhindered by our knowledge (or lack of knowledge) of theirtaxonomic status (Daugherty et al. 1990, May 1990,
O’Brien and Mayr 199 1). Taxonomic questions involving species, subspecies, hybrids, and inbreeding effects
will become increasingly important as wild populations
become smaller and increasingly isolated and as captive
populations are managed more intensively.
The taxonomy of the cat species is an area of much disagreement, as previously discussed by Lars Werdelin. For

into a number of different genera, just as has been done inother families of carnivores. The view of the Felidae as
including only the genera Felis, Panthera, and Acinonyx isonly seen in the non-specialist literature and should belaid to rest once and for all.

Molecular Genetics andPhylogeneticsof the Felidae
by Stephen J. O’Brien
A fundamental component of conservation strategies forthreatened species is the systematic classification of
species and significantly differentiated populations below
the species level (subspecies). Uncertainty over the units

Evolution-------

l

-----

I
v---l
I

I

(4

I

r ---I-

I

I

I

I

15

10
years before

5MillionFigure 3. Phylogenetic(S.J. O’Brien).

-of the Felidaeof the Felidae

lionjaguarPant herasnow leopard
genus
leopardtiger
cloudedleopardmarbled cat
bobcatIberian/Pardel
lynx
Eurasianlynx
Canadianlynxjaguarundi
caracalservalfishing cat
Pantherarusty-spottedcatlineageleopard cat
flat-headedcatAfrican golden cat
Borneanbay catTemminck’s
golden cat
cheetahpuma, mountainlion, cougar
Pallas’s cat
Asian steppe cat
Domestic
cat
Chinesemountaincatjungle cat
lineagesand cat
black-footed
cat
Europeanwild cat
African wild cat
domestic
cattiger cat, oncillakodkod, guina, huina
OcelotGeoff roy’s cat
lineagepampas cat

Pantheralea
Panthera
onca
Panthera
uncla
Panthera
pardusPan thera tlgris
Neofells
nebulosa
Pardofelis
marmorata
Lynx &us
Lynx pardinus
Lynx lynx
Lynx canadensls
Herpallurusyaguarondl
CaracalcaracalLeptailurus
servalPrionailurus
viverrinus
Prlonailurusrubiginosus
Prionallurus
bengalensis
Prionailurus
planlceps
Profelis
aurataCatopumabadia
Catopuma
temmlncki
A&onyxjuba tus
Puma concolor
Otocolobus
manul
Felis silves tris orna ta
Fells bletl
Fells chaus
Fells margarita
Fe/is nlgrlpes
Felis silvestris
Fe/is silves tris lib yea
Fe/is catus
Leopardus
tigrlnus
Oncifelis
guignaOncifelisgeoffroyl
Oncifelis
colocolo
Leopardus
wledii
Leopardus
pardalls

present

relationshipof felid species

basedon a consensus

.*.

XXIII

of molecular,

karyologic,

and morphologicalcharacters.

1

Taxonomyof the Felidae

example, the most recent edition of Waker’s Mammals of
the World (Nowak and Paradiso 1991) lists four different
taxonomic schemes (after Leyhausen, Hemmer, Ewer, andothers), which lump cat species into as few as four generaor split them into as many as 19.
Paleontologists tell us that the two carnivore families,
Felidae and Canidae, diverged from a common ancestor
about 50 million years ago because the “missing link” fossils that share characteristicsof the two families areapproximately this age. By measuring the quantitative differences that occu r in genes and DNA sequence s of catsand dogs, we have developed a good qu antitative estimateof the amount of mutational change that occurred in thesegroups over the past 40 million years. This calibration,termed the “molecular clock,” is not the perfect evolutionary timepiece, but it has helped resolve a number of controversies in evolutionary studies.
Several molecular metrics have been applied to estimate relationships between cat species, using blood andskin cell cultures as the biological materials. The Felidae
is a relatively difficult group to analyze in this way, as
there are many species which have split from each otherrelatively recently. Five different methods have been
applied to samples from living cat species. Three of these,
allozyme genetic distance (O’Brien et al. 1987d), 2DE
genetic distance (Goldman and O’Brien 1993), and albumin immunological distance (Collier and O’Brien 1985),
measure differences in protein (gene product) sequences.
Two methods, DNA-DNA
hybridization (Wayne et al.
1989) and DNA sequence analysis, compare the specific
DNA sequence code of different cat species.
The results are neither perfect nor camp lete, but they
have converged on several conclusions and on a “best”
phylogenetic tree based on concordance of the various
molecular tests. The molecules, when calibrated and interpreted along wi th certain fossil . remains, describe a scenario that is summarized in Fig. 3. The major conclusion
derived from the molecular topology was the resolutionof felid evolution into three major lineages. The earliest
branch occurred approximately 12 million years ago andled to the small South American cats (ocelot, margay,
oncilla, Geoffroy’s cat, and others). The second branchingoccurred about 8 to 10 million years ago and included theclose relatives of the domestic cat (wildcats, jungle cat,sand cat, black-footed cat) and the manul. About 4-6 million years ago a gradual divergence of mid-sized and largecats began: the most recent (1.8-3.8 million years ago) produced a split of the lynxes and the big cats.
One dramatic surprise revealed by the molecularmethod was the placement of the morphologically specialized cheetah in the midst of the mid-sized cat radiation. Earlier taxonomists had largely agreed that the
cheetah’s adaptive specializations for high-speed sprinting merited separate generic status and likely indicatedan early divergence from the felid evolutionary tree. The
molecules did not agree. In addition, recent DNA
sequence data on mitochondrial DNA genes suggest thatthe cheetah’s closest living relative is the American puma
(Janczewski 1993).
Re-examination of other non-molecular characters of
the Felidae in the context of the molecular trees has not
only reinforced certain patterns, but has also shed light on
the evolutionary processes that occurred in this group. For
example, the chromosomes of all the major cat groups(that is, big cats, domestic cat relatives, and SouthAmerican small cats) look identical within the clusters, but
distinct from other groups. Further, many of the anatomical similarities between cat species that have confounded
the experts are now beginning to make more sense. We
certainly do not have all the answers yet, but the recent
advances in our understanding of molecular evolution ofcat genomes offers the prospect that resolution of thesethorny taxonomic issues may now be within our reach.
Properly interpreted, a consensus molecular, morphological, and ethological classification scheme would provide
a framework for conservation programs.

xxiv

Part I: Species

Accounts.

introduction

Part I
Species Accounts
Introduction

Common names of prey species are used, and their scientific names listed in Appendix 3.

The cats are grouped according to the five geopoliticalregions in which they occur: (1) Sub-Saharan Africa; (2)North Africa and Southwest Asia; (3) Tropical Asia; (4)Eurasia; and (5) the Americas. There are no cats (otherthan domestic) in Australasia and Oceania.
Some cats occur in more than one region. Where there
is sufficient information, an account has been written foreach region in which a species occurs (cheetah, caracal,
wildcat, lion, and leopard); otherwise, a single species
account is included under the region with which thespecies is most strongly associated.
Each regional chapter opens with a table which ranks
the vulnerability of the species occurring in the region.
Species Accounts are presented in that order. This introduction explains the structure of the Species Accounts andthe ranking of species vulnerability.

Biology
This section includes basic biological data, which are generally sparse, and typically derived from captive animals(labelled C in the Species Accounts).
Information
obtained from studies of cats in the wild (labelled W) is
often known for only a small portion of the total range.
Populations elsewhere may differ significantly (e.g. seasonality of reproduction, longevity, mortality rates).
Habitat and Distribution
Habitat preference and association is discussed, and distribution is illustrated in range maps.
Population Status
Vulnerability ranking (see following section for explanation) and status according to the 1994 IUCN Red List of
Threatened Animals (Groombridge 1993: see Box 2) are
given, and current information on the status of wild populations is presented, including data on density and homerange size, where available.

Structure of theSpecies Accounts
Other Names
Species names are given in local languages within theirrange, as well as in the three international languages:
French, German, and Spanish. Readers are invited to forward other local names, or correct any given in the Species
Accounts, in order to build up the database.

Protection Status
International protection: all cats were listed on either
Appendix I or II of the Convention on International Tradein Endangered Species of Wild Fauna and Flora (CITES)
by 1977, so that international commerce in cats (dead or
alive), their pelts, and other products has been either prohibited or regulated since that time (see the Trade chapter
in Part II). Dates are given for cases where CITES listings were subsequently changed. National legislation: categorizes the type of legal protection cats receive in theirrange states.

Description and Behavior
Because photographs are provided, physical descriptions
are kept to a minimum, but include general appearance,
distinguishing features, and adult weight. Readers shouldbear in mind that body weight can be substantially influenced if the cat has a full stomach: for example, Wilson(1968) reported that the stomach contents of a 43 kg
female leopard weighed 6.6 kg, or 18% of her bodyweight. It was not generally possible to distinguish fromthe literature whether reported weights accounted forstomach contents. Characteristic aspects of the species’
behavior and ecology, including diet, are discussed.

Occurrence in Protected Areas
Protected areas where the species is known or suspected tooccur are shown on the distribution maps. Information on
occurrence in protected areas was gathered from a wide

1

Part I: Species

Accounts.

Introduction

variety of sources, including IUCN protected area directories (IUCN 1982, 1987a, 1990a, Green 1993-withreported occurrence independently confirmed where possible), the voluminous files of the Protected Areas Data
Unit of the World Conservation Monitoring Centre in
Cambridge, databases maintained by national governments and institutions, the literature and, most importantly,
data provided by correspondents. Generally, priority was
given to larger reserves, but the data are patchy and themaps reflect this. For some species, it was not possible todisplay all protected areas in which presence is known;
for others, occurrence is insufficiently known and only afew protected areas are indicated. For a very few species,
the amount of habitat needed to support minimum viable
populations has been calculated, and protected areas of
the requisite size are marked with a square on the maps.
Protected areas are named according to the management categories developed by the IUCN Commission on
National Parks and Protected Areas, which are used by
the United Nations in their List of National Parks andProtected Areas (IUCN 1990b), and by the Protected
Areas Data Unit of the World Conservation Monitoring
Centre, which maintains an extensive international database. The categories standardize the type of legal protection and management extended to an area.
I.
II.
III.
IV.
V.
VI.
VII.
VIII.
IX.

CategorizationofSpecies Vulnerability
A system to rank species according to their vulnerability to
extinction was developed for this Action Plan. There are
five categories of vulnerability, with “1” the highest.
Species are ranked on a global scale (in relation to all othercat species) in Box 1, as well as a regional scale (in relation
to other cat species occurring there). Regional rankings
are summarized at the start of each regional chapter in PartI, and both global and regional species rankings are givenin the Species Accounts under Population Status.
The ranking system was developed in order to provide
an objective method for prioritizing species and populations for conservation.
IUCN maintains a Red List of
species of conservation concern, but the criteria for theircategories of threat were not quantitative, and new criteria have been developed (IUCN Species Survival
Commission 1995). Because many people are accustomed
to the old system, and for purposes of comparison, the1994 Red List rankings are also given in the Species
Accounts. Definitions of the IUCN rankings are given in
Box 2.
Most attempts to rank species vulnerability objectively,
including the new IUCN threat criteria, involve estimatesof population size and/or rate of decline. However, given
the paucity of data on density and species presence orabsence, it is not possible to derive reliable quantitative
estimates of total numbers or rate of change in abundancefor cats (see Part II, Chapter 3 for discussion of the difficulties of counting cats). The method used in this document to rank species vulnerability is based on other factors
which influence population size and extinction risk: habitat
association, geographic range area, and body size. Huntingpressure is also accounted for as an active threat with thepotential to remove animals from otherwise viable portionsof their range. For global comparison, each cat species wasscored for these criteria as described below. For regional
rankings, the criteria are the same but the scoring may differ (see the introductions to each regional chapter).

Scientific Reserve/Strict Nature Reserve
National Park
Natural Monument/Natural Landmark
Managed Nature Reserve/Wildlife Sanctuary
Protected Landscape (recreational activities
predominate)
Resource Reserve
Natural Biotic Area/Anthropological Reserve
Multiple-Use Management Area/Managed
Resource Area
Biosphere Reserve

If a category has not been assigned, the full name of thereserve is given. Protected areas which have been listed as
Biosphere Reserves under the UNESCO Man and the
Biosphere program are designated *. Protected areas
which have been accepted as World Heritage Sites under
the Convention Concerning the Protection of the World
Cultural and Natural Heritage ( 1975) are designated **.
Protected areas which qualify as both are denoted #.

Criterion 1. Habitat Association
Species which are associated with a narrow spectrum of
habitats are more vulnerable to extinction than species
which are more broadlv associated.

Principal ThreatsA brief overview of the major threats, focusing on thosethat particularly affect that species. Threats affecting catsin general are discussed at length in Part II, rather than in
the Species Accounts.

The occurrence of cat species in a standard set of global
habitat types (Olson et al. 1983) was evaluated. The habitat classification is described in Part II, Chapter 1, and mapsof the global distribution of these habitat types are included.
The degree of species association with a particular habitat
type was assigned as strong, significant, marginal, or

Action Planning
A link to relevant priority projects in Part III.

2

Pat? I: Species

Box 1
Worksheet Summary for Global Cat Species Vulnerability

Accounts.

Introduction

Score

Total
Score

Rankings

Habitat Association
St [h/Jar](Tot) Score

Geog.
Score
Range
(106km2)

Body
Sine
in kg

N: 3

[3]

(6)

-1

R:

O-08

-2

M:

9.3

0

-3

I: 6
I: 1
N: 2
N: 2
N: 3
N: 2
N: 2
N: 3
N: 5
N: 3

-31 (9)61 (7)PI (2)
31 (5)PI (3)
21 (4)PI (2)

0
0
-1
-1
-1
-1
-1

71 (5) -1

s:
S:
R:
R:
R:
R:
R:
R:
S:
S:

1.99
2.39
0.05
0.29
0.95
0.16
0.62
1.18
2.33
2.46

-1
-1
-2
-2
-2
-2
-2
-2
-1
-1

L:136.0
1: 37.5
S: 2.4
S: 6.5s: 1.2s: 2.2
5: 4.0s: 1.9
M: 6.8
M: 10.0

-1
-1
+I
+I
+I
+I
+I
+I
0
0

-2
-2
-2
-2
-2
-2
-2
-2
-2
-2

I: 4
I: 5
I: 4
I: 5
N: 3
I: 7
I: 4
N: 3

[4]
[2]
[3]
[3]
[I]
[O]
[4]
[I]

(8)
(7)
(7)
(8)
(4)
(7)
(8)
(4)

0
0
0
0
-1
0
0
-1

M:
M:
M:
S:
S:
R:
S:
S:

7.35
7.18
8.91
2.66
2.90
0.78
2.79
2.42

0
0
0
-1
-1
-2
-1
-1

L: 43.0
1: 126.0
1: 56.0
M: 10.0
s: 2.0s: 1.5
M: 20.0
s: 3.5

-1
-1
-1
0
+I
+I
0
+I

-1
-1
-1
-1
-1
-1
-1
-1

N: 2
N: 2
I: 7
I: 4
I: 6
N: 4

[I]
[3]
[2]
[4]
[I]
[2]

(3)
(5)
(9)
(8)
(7)
(6)

-1
-1
0
0
0
-1

M:
M:
M:
M:
S:
M:

5.40
6.06
8.18
5.06
2.80
5.08

0
0
0
0
-1
0

S: 2.5
5: 3.2
M: 10.0
M: 8.5
S: 4.2
5: 3.0

+I
+I
0
0
+I
+I

0
0
0
0
0
0

Pampas cat, 0. colocolo

B: 8
B:lO
I: 5
B: 6
B: 7
B: 4

[7]
[5]
[4]
[S]
[4]
[6]

(15)
(15)
( 9)
(12)
(11)
(10)

+I
+I
0
+I
+I
+I

W: 17.12
W: 23.14
W: 12.45
W: 13.56
M: 7.24
S: 3.86

+I
+I
+I
+I
0
-1

1: 41.0
1: 40.0
M: 8.8
M: 17.0
M: 7.5s: 3.4

+I
-1
0
0
0
+I

+I
+I
+I
+I
+I
+I

Category 5b
Caracal, C. caracal
Jungle cat, F. chaus
Leopard cat, P. bengalensis

B: 6
B: 8
B: 7

[4] (10)[5] (13)[S] (12)+I+I+I

W: 18.99
M: 8.49
M: 8.66+I00

M: 10.0
s: 5.4
S: 2.4

0
+I
+I

+2
+2
+2

SpeciesCategory 1
lberian lynx, L. pardi~~us

Category 2
Tiger (A), P. tigris
Snow leopard (A), U, uncia
Bornean bay cat, C. badia
Chinese mtn. cat, F. bieti
Black-footed cat, F. nigripes
Kodkod, 0. guigna
Andean mtn cat, 0. jacubitus
Flat-headed cat, P. planiceps
Fishing cat, P, viverrinus
African golden cat, P. aurata

‘01
;I]

(3)
(6)

-1
-1

Category 3
Cheetah (A), A. jubatus
Lion (A), P. lea
Jaguar (A), P. onca
Asiatic golden cat, C. temmincki
Oncilla, L. tigrinus
Rusty-spotted cat, P. rubiginosus
Clouded leopard, iV. nebulosa
Marbled cat, P. marmorata

Category 4
Sand cat, F. margarita
Margay, L. wiedi
Serval, L. serval
Canada lynx, L. canadensis
Geoffroy’s cat, 0. geoffroyi
Manul, 0. manul

Category 5
Category 5a
Puma (A), P. concolor
Leopard (A), P. pardus
Ocelot, L. pardalis
Eurasian lynx, L. /ynx
Bobcat, L. rufus

Continued on next page3

Part I: Species

Accounts.

Introduction

Species

Habitat Association
St [Mar] (Tot) Score

Geog.
Range
(106 km*)

Score

B: 8
B: 6

w: 34.17
w: 13.53

+I
+I

Body
Size
in kg

Score

Total
Score

+I
+I

+3
+3

Category 5c
Wildcat, F. silvestris
Jaguarundi, H. yaguarondi[43 (12)[4] (10)

+I
+I

s:
s:

3.5
4.4

Key:
Habitat Association
St = Number of strong + significant habitats
N = Narrow; I = Intermediate; B = Broad[Mar] = Number of marginal habitats(Tot) = Total number of habitats

Geographic Range
R = Restricted (c 1.5 million km*)
S = Small (1.6 - 4 million km2)
M = Medium (5-9 million km2)
W = Wide (1 O-35 million km2)

6ody Size
L = Large; M = Medium;

S = Small(A) = Actively threatened

absent. For example: the sandcat is strongly associatedwith sandydesert;the lion is significantly associatedwith
grasslandand shrubland; the snow leopard is marginallyassociatedwith coniferousforest; andthe Andeanmountaincat is absentfrom broad-leavedhumid forest.
The Habitat chapter describeshow degreeof association was determined,and Appendix 4 lists habitat associationsfor each specieson both a global andregional level.
For vulnerability ranking, specieswere scoredasnarrow,intermediate or broad in habitat associationon the basisof the total number of habitat types in which a species
occurs (strong, significant, or marginal).
Narrow habitat association(N): 2-6 habitat types(14 species).Score: -1.
Intermediatehabitat association(I): 7-9 habitat types(12 species).Score: 0.
Broad habitat association(B): lo-15 habitat types(10 species).Score: +l.

Criterion 2. Geographic Range Size
Specieswith a restricted geographic range are more vulnerable to extinction than specieswith a wide range.
Range size was calculated (in millions of km2) by comparing the range maps(SpeciesAccounts) to the global
habitat maps(Habitat chapter), applying reduction factors
asnecessary(for occurrencein only part of a given habitat type), andaddingup the geographicareafor eachhabitat type asderived from Olson et al. (1983). Only strongor
significant habitat associationswere used;habitatsclassified as marginal for a specieswere not included in thecomputation of its geographicrange size. The methodology is describedin greater detail in Appendix 4.
This exercise was undertaken only to derive a basis,
more objective than a visual examination of distribution
maps,for comparing speciesrange size. However, for avariety of reasons the potential for error is high (seeAppendix 4), and the figures given shouldnot be treatedasdefinitive. They appearin the worksheet summary of
global cat speciesvulnerability rankings(Box 1) for com-

Pat? I: Species

Accounts.

Introduction

parative purposes, but are not given in the species accounts.
Restricted geographic range (R): <I.5 million km2(8 species). Score: -2.
Small geographic range (S): 1.6-4 million km2(10 species). Score: -1.
Medium geographic range (M): 5-9 million km2(9 species).Score: 0.
Wide geographicrange (W): lo-35 million km2(8 species).Score: +I.

Criterion 3. Body Size
The larger an animal, the larger its homerange, the lower
its density, and the greater its local rarity.

Box 21994 IUCN Threatened Species
categories(Groombridge

1993)

Extinct
Species not definitely
past 50 years.

locatedin the wild during theEndangered
Taxa in danger of extinction and whose survival isunlikely if the causal factors continue operating.
Included
are taxa whose numbers
have been
reduced to a critical level or whose habitats havebeen so drastically reduced that they are deemed
to be in immediate danger of extinction.

Vulnerable
A number of studies have found a general relationshipbetween body size and density (Harestad and Bunnell
1979, Eisenberg1980, Arita et al. 1990). Within a givenarea(suchasa nature reserve),big catsare expected to be
morerare than smallones. Body size is thusa usefulindex
to relative abundance(Soule 1991), and it was included
asa criterion to provide a link to speciespopulation size
estimates,which are frequently requested. Average adultfemale weight was usedto rank speciesin all but a few
cases,when average weight (no gender specified) was
used:thesespecieswere all smallcatsunder 7 kg.
Large body size (L): 35-135 kg (7 species).Score: -1.
Medium body size (M): 7-20 kg (11 species).Score: 0.
Small body size (S): <6.5 kg (18 species).Score: +I.

Criterion 4. Active Threat
Widespread and uncontrolled hunting, which has thepotential to remove animalsfrom viable habitat in which
they would otherwise be present, is an active threat
to species.
Habitat lossand change are gradual, ongoing processes
affecting all species,but widespread and uncontrolled
hunting, which may be for food, sport,or trade, is an active
threat to both catsandtheir prey. It is relevant primarily tothe big cats. Their prey are mainly large ungulates,which
aremore vulnerableto over-hunting than rodentsand othersmall mammals, on which small cats mainly subsist.
Moreover, big cats are more likely to be commerciallyhuntedandpersecutedasproblemanimals. The suffix “A”
is appendedto the vulnerability rankings of all the larger

Taxa believed likely to move into the “Endangered”
category in the near future if the causal factors continue operating.
Included are taxa of which most or
all of the populations
are decreasing
because ofoverexploitation,
extensive destruction of habitat orother environmental
disturbance; taxa with populations that have been seriously depleted and whose
ultimate security has not yet been assured; andtaxa with populations that are still abundant but areunder threat from severe adverse factors throughout
their range.
/MB. In practice, “Endangered”
and “Vulnerable”
categories may include, temporarily, taxa whose populations are beginning to recover as a result of remedial action, but whose recovery is insufficient
to justify their transfer to another category.

Rare
Taxa with small world populations
that are notat present “Endangered”
or “Vulnerable,”but areat risk. These taxa are usually localized within
restricted geographical areas or habitats or are thinly
scattered over a more extensive range.

Indeterminate
Taxa known to be “Endangered,”“Vulnerable,”or“Rare,” but where there is not enough information
to say which of the three categories is appropriate.

lnsuff iciently Known
Taxa that are suspec&d but not definitely known tobelong to any of the above categories, because oflack of information.

Part I: Species

Accounts.

Introduction

cats because of the relatively high levels of hunting pressure facing these species.

a category, actively threatened species (“A”) are listed first
because of their greater vulnerability. The more common
and less vulnerable species (scores +l to +3) were grouped
under Category 5 (5a, 5b, 5~). This category is of lowest
conservation priority on a global scale. However, regional
rankings may differ. A summary of species vulnerability
on a regional scale precedes each regional chapter of theSpecies Accounts.

Scoring
The worksheet summary (Box 1) presents the scoring andvulnerability rankings of species on a global level. The
higher the ranking, the higher the extinction risk. Within

6

Pat? I: Species

Accounts.

Chapter

1. Sub-SaharanPart I
Species Accounts

Chapter 1
Sub-Saharan Africa

Box 1
Vulnerability

Index to Species of the Region (in order of vulnerability)

Species

Habitat Association
St [Mar] (Tot) Score

Geog.
Range
(106 km*)

Score

Black-footed cat, F. nigripes*
African golden cat, P. aurata*
Cheetah, A. jubatus*
Lion, P. lea*Set-vat, L. serval*
Leopard, P. par&s
Caracal, C. caracal*
African wildcat, F. s. lyhica group

N: 3 [O]N: 3 [Z]1:4[4]

R: 0.95
S: 2.46
M: 6.33
M: 7.15
M: 7.91
W: 14,56
w: 11.93
w: 16.80

-2
-1

(3)
(5)
(8)
(7)
1:5[2]
1:6[2](8)
B: 7 [3] (IO)
l:5[4](9)
I:6[2]
(8)

-1
-1
0
0
0
+I
0
0

Body

Size
Score

0
0
0
+I
+I
+I

s +I
M 0
L -1
L -1
M 0
L -1
MO
s +I

Total
Score

Ranking

-2
-2

1
1
2(A)
2(A)
3
4(A)
4
5

-1

-1
0
+I
+I
+2

Key:
*Most or all of this species’ range lies within the region

Habitat Association
St = number of strong + significant habitats
N = Narrow (-1); I = Intermediate (0); B = Broad (+I)
[Mar] = number of marginal habitats(Tot) = total number of habitats

Geographic Range (in millions of km*)
R = Restricted (-2); S = Small (-1);

M = Medium(0); W = Wide (+I)

Body Size
L = Large (-1); M = Medium(0); S = Small (+I)

(A) = Actively threatened

L

Regional Criteria
Habitat Association: Narrow = 3-5 habitat types; Intermediate = 7-9 habitat types; Broad = IO habitat types
Geographic Range: Restricted = cl million km*; Small = I-6 million km*; Medium = 6-9 million km*;
Wide = 9-I 7 million km*
Body Size: Large = 35-I 35 kg; Medium = 7-20 kg; Small = ~6.5 kg

See the Introduction

to the Species

Accounts

for explanationof the vulnerability

7

rankingsystem(pp. 2-6).

AfricaPart I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Black-footed

catBlack-footed cat, Fe/isnigripes Burchell, 1824

disused burrows, such as those made by springhares, or inrocky crevices (Shortridge 1934, Sliwa 1993). Blackfooted cats are apparently water-independent (Skinner andSmithers 1990). Unusually high blood levels of creatinine and urea (even for arid-adapted cats) have been foundin both wild and captive black-footed cats (G. Olbricht and
A. Sliwa, pers. comm. 1993). Olbricht and Sliwa have also
noted that black-footed cats appear to have higher energy
requirements than the larger African wildcats.

Other Names
Small spotted cat (English); chat a pieds noirs (French);
Schwarzfusskatze (German); gato patinegro, gato de pies
negros (Spanish); klein gekolde kat, swart poot kat, miershooptier [anthill tiger] (Afrikaans: South Africa); !koirus(Nama: Namibia); tutchu (Naron Bushman: Botswana);
sebala, lototsi (Setswana: Botswana); ingwe yeziduli(Xhosa: South Africa).

Biology
Birth season: (W) a pregnant female carrying two fetuses
was collected in South Africa’s Transvaal province inNovember (Rautenbach 1978). A kitten approximately
one month old was observed in January in the northern
Cape (A. Sliwa in litt. 1993); and two kittens were born inlate February in a den in a hollow termite mound in thesame area (A. Sliwa in litt. 1994).

Description
and Behavior (Plate 3)
The black-footed cat is among the world’s smallest felines,
with females weighing around 1.2 kg (range 0.8 1.6) andmales larger at 1.6-2.1 kg (Smithers 1971, Stuart 1981,
Lynch 1983, A. Sliwa in litt. 1993). Total length ranges
from 50-63 cm (Smithers 1971), and shoulder height isaround 25 cm (Stuart and Wilson 1988). It is boldly patterned with blackish oblong spots, and its legs are barredwith thick dark stripes. The undersides of its feet areblack, like those of the African wildcat. The auditory bullae are enlarged, with total length about 25% of skull
length (Skinner and Smithers 1990).
The diet consists mainly of small mammals and birds,
and also includes arachnids, insects, and reptiles (Rautenbath 1978, Smithers 1971, Stuart 198 1, Sliwa 1994).
Radio-collared cats were observed by Sliwa (1994) tocatch larks by stalking to within a short range and making
a quick run and jump, catching some in the air as they flew
off. Small rodents were caught by stalking or waiting (upto 30 min.) at holes. They also fed on emerging alates of
the harvester termite, and caught larger winged insects
such as grasshoppers. The largest mammal prey was anadult Cape hare, weighing as much as the adult female
who caught it (1.5 kg). The largest bird caught was a smallbustard, the black koorhaan, weighing 700 g. Blackfooted cats have also been observed to eat black koorhaan
eggs: “She flushed a female koorhaan from her nest, andthen crushed the eggs gently between her jaws and licked
their contents clean” (A. Sliwa in litt. 1994). Stuart (198 1)
reports a black-footed cat trapped with a guinea fowl carcass as bait. A. Sliwa (pers. comm.) has observed blackfooted cats caching rodent and bird carcasses in hollows,
returning after 2-14 hours to feed, and once observed acat scavenging for four nights on a springbok lamb.
Most observations in the wild have been at night(Smithers 1971; P. Stander, J. Visser, pers. comm.). The
cat observed by Sliwa (1993) was generally active
between sunset and sunrise, and only during the cold winter months at first light and in late afternoon. The cat was
active for most of the night, travelling an average of 8 km
while foraging (n=lO nights). Black-footed cats lie up in

Estrus: (C) l-5 days (Leyhausen and Tonkin 1966, Mellen1989) .
Estrus cycle: (C) 54 days (Mellen 1989).
Gestation: (C) 63-68 days (Leyhausen and Tonkin 1966).
Litter size: (C) 1.7 1 t 0.18 (n=9: Mellen 1989); range l2, rarely 3 (Visser 1977, Armstrong 1978).
Age at sexual maturity: (C) females 12 (Mellen 1989) -21
months (Leyhausen and Tonkin 1966); onset of spermatogenesis in males at about one year (R. Evans in litt. 1993).
Longevity:- (C) up to 13 years (Green 1991).
Habitat and Distribution
The black-footed cat is restricted to the arid lands of southern Africa (Fig. 1). It is typically associated with open,
sandy, grassy habitats with sparse shrub and tree cover,such as the Kalahari and Karoo regions (Smithers 197 1,
1975; Visser 1977, Mills et al. 1984, Stuart and Wilson
1988, Sliwa 1993). A. Sliwa (pers. comm.) describes longgrass with high rodent and bird densities as optimal habitat.
The northernmost records are from around 19” S in
Namibia and Botswana (Shortridge 1934, Visser 1978, P.
Stander, pers. comm. 1992), although the species may
occur in the southwestern comer of Angola (Anstey 1992).
It has not been recorded from Zimbabwe and Mozambique, although it probably occurs there marginally(Shortridge 1934, Dias 1966, Stuart and Wilson 1988, J.
Visser in Zitt. 1993). A record for Malawi is erroneous(Ansell and Dowsett 1988).
Population Status
Global: Category 2. Regional: Category 1. IUCN: notlisted. Most authorities have described the black-footed
cat as a naturally rare species (Stuart and Wilson 1988,

8

Part I: SpecieskniSkinner and Smithers 1990). Shortridge (1934) reported
at the turn of the century that mantles made from the skinsof this species were expensive, “on account of theirscarcity.” Still, it is locally common at certain localities inSouth Africa, especially in the Orange Free State andnorthern Cape (J. Visser in Zitt. 1993). Being restricted toarid environments, it probably occurs at relatively low
densities. An adult female observed for three months ona game farm near Kimberley (northern Cape, South
Africa) had a home range of 12 km? A young maleobserved for a shorter period of time maintained a homerange of 13 km2, overlapping the female’s range by about
50% (A. Sliwa in litt. 1993).
Protection Status
CITES Appendix I. National legislation: protected acrossmost of its range. Hunting prohibited: Botswana, South

Accounts.

Chapter

1. Sub-SaharanAfrica,Black-footed

cat

Africa. No legal protection: Mozambique, Namibia,
Zimbabwe (IUCN Environmental Law Centre 1986; P.
Norton, C. Stuart in litt. 1993).
Principal Threats
Indiscriminate methods of predator control could be a significant threat, although farmers seldom report capturing
black-footed cats in problem animal surveys (Joubert et al.
1982, Stuart and Wilson 1988). Farmers in South Africa
and Namibia consider the similar-looking African wildcat
a predator of small livestock, and set out steel-jaw traps
and poisoned bait to get rid of them (Joubert et al. 1982,
Vorster 1988). Carcass poisoning for jackal control could
be a threat to the black-footed cat, which readily scavenges(A. Sliwa, pers. comm.). A similar threat is poisoning of
locusts, which are food for the black-footed cat. Finally,
overgrazing by livestock is prevalent throughout the

Figure 1. Distributionof theblack-footedcat (F. nigripes).
I. Etosha II (Namibia);
2. Hwange II complex(Zimbabwe);
3. Makgadikgadi
Pans IV (Botswana);
4. Gemsbok II (Botswana)+ Kalahari
Gemsbok
II (South Africa)
complex; 5. Karoo II; 6. Addo
Elephant II; 7. BarberspanIV;8. specimen
collected at Marble
Hall, Transvaal
province (Skinnerand Smithers 1990); 9. Kittencollected in northwesternNatalprovince (Rowe-Rowe1992);
IO. Royal Natal II; 11. Loteni IV;
12. Willem Pretorius
IV (South
Africa); 13. lona VI (Angola).

Par? I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Africangoldencat

species’ range, and habitat deterioration can lead to reductions of the cat’s small vertebrate prey base (A. Sliwa,
pers. comm.).

in trees to some extent (Basilio 1962, Kingdon 1977), but
probably catches most of its prey on the ground (J. Hart
and M. Katembo in prep.). Hart and Katembo analyzed 60
golden cat scats from Zaire’s Ituri Forest, and found that
5 1% contained rodents and 20% ungulates. The rodents
were mostly small species weighing less than 300 g. From
carcass collections, they also note that scavenged eagle
kills and predation on fallen, injured primates may be an
important component of rain forest felid diets.
Hart and Katembo’s data serve to balance anecdotalreports that golden cats prey mainly on small to mid-sized
mammals, including tree hyraxes, the larger rodents(Basilio 1962, Brooks 1962, Rahm and Christiaensen
1963), and smaller forest antelopes (Van Saceghem 1942,
Carpaneto and Germi 1989). On the contrary, they foundsmall rodents to be more important. Other data on diet are
patchy. For example, the stomach of one golden cat from
Senegal contained the remains of a bird (Gaillard 1969),
and Kingdon (1977) found the remains of red duikers,
monkeys, rodents, and birds in scats examined from
Uganda’s Bwindi National Park. D. Jenny (pers. comm.)
found many pangolin remains in scats in Tai’ NP, Ivory
Coast. M. Agnanga (in lift. 1993) includes fish in the diet.
Although there have been reports of predation on domestic animals, including chickens, goats, and sheep(Gyldenstolpe 1928, Bourdelle and Babault 193 1, Kingdon
1977), such predation appears to be rather rare (E. Abe,M. Agnanga, B. Hoppe-Dominik, S. Lahm in Zitt. 1993).

Action Planning
Projects 18 and 19.

African golden cat, Profelis
aurata (Temminck, 1827)
Other Names
Chat dare africain (French); Afrikanische
Goldkatze(German); gato dorado (Spanish); gnaou ya zamba(Lingala: west Africa); lobwa, ebyo, ebie (Kota, Fang,
Kwele: Gabon); embaka, ekinyange, semaguruet (Lukiga,
Lukonjo, Kipsigi: east Africa); soukalan (Mandinka);osolimi, makolili, akalwa, egabasoti, esele, a’ka (Mbuti
Pygmies: Zaire); donnou, dondou (Peul).
Description
and Behavior (Plate 1)
The African golden cat is a medium-sized cat. Adult
males weigh 11-14 kg (Van Mensch and Van Bree 1969,
S. Lahm in Zitt. 1993). The only recorded weight of a wild
female is 6.2 kg (Van Mensch and Van Bree 1969), but
this was probably an immature animal. The African
golden cat has both a reddish-brownand greyish color
form, and its coat can be spotted or plain. Pocock (1907a)
described an animal in the London Zoo whose color
changed entirely from rufous to grey in four months. Van
Mensch and Van Bree (1969) examined 186 pelts fromvarious localities and found that 50% were of the red phaseand 46% of the greyish phase, with 4% being totally black.
While color phase appears variable across its range, theyfound that specimens taken from west Africa tended to be
more spotted than those from east-central Africa, with theZaire River forming an approximate boundary. The white
underbelly is consistently marked with large black spots.
Despite a striking external similarity to the Asian
golden cat, many authorities believe that the two species
are not closely related (Van Mensch and Van Bree 1969,
Hemmer 1978a, Wozencraft 1993; but see Fig. 3 underTaxonomy). The similarity of the golden cats may haveresulted from convergent evolution in moist forest habitat, as there has been no direct forest connection between
Africa and Asia for 20 million years (Groves 1982), but
the relationship still deserves closer examination.
The African golden cat has never been studied and little
is known of its behavior. It is reported to be primarily nocturnal and to rest in trees during the day (Rosevear 1974,
Guggisberg 1975, Kingdon 1977, Happold 1987). Diurnal
activity has also been noted (Kingdon 1977). It may hunt

Biology
Litter size: (W) According to the Mbuti Pygmies of northeastern Zaire, one (Carpaneto and Germi 1989). J. Hart
and M. Katembo (in prep.) also found one nursing kitten ina fallen, hollow log. No other information.
Habitat and Distribution
The primary habitat of the African golden cat is the moist
forest zone of equatorial Africa, including mangrove andalpine bamboo forests. Golden cats can penetrate savannah grasslands along belts of riverine forest (Van Menschand Van Bree 1969), and so their distribution probably
extends beyond the moist forest zone. As an extreme
example, the species was recorded from Nioro du Sahel,Mali, in relatively arid Savannah woodland (Bigourdan andPrunier 1937), although possibly in error (Van Mensch and
Van Bree 1969).
Golden cats apparently adapt well to logged areas, as
destruction of the canopy favors the dense secondary
undergrowthwith which they are often associated(Kingdon 1977, Anstey 1991, S. Lahm in Zitt. 1993). Edgeenvironments generally contain higher rodent densities,
and may thus be preferred (J. Hart in Zitt. 1994). However,primary forest with minimal human disturbance is the10

Part I: Species

Minimalty disturbed
tropical rainforest

Cl
cl
.

Protected area where
species occurs#

Protected area where
species may occur

primarynUnconfirmed

Chapter

1. Sub-SaharanAfrica,Africangoldencat

Figure 2. Distributionof theAfrican golden cat (P. aurata).
1. Record from Nioro du Sahel,
Mali (Bigourdan
and Prunier1937); 2. Basse-CasamanceII(Senegal);
3. Gola Forest
Reserve(Sierra Leone); 4. Mt
Nimba I# complex (Ivory Coastand Guinea); 5. Sapo II (Liberia);
6. Tai’ II# complex;
7. Mt. Sangbe
and Mt. Peko II; 8. Comoe II#(Ivory Coast); 9. Bia II (Ghana);
IO. Gashaka-Gumti
II (Nigeria);
11. Korup II; 12. Dja IV# (Cameroon); 13. Lope IV (Gabon);
14. Odzala II* complex (Congo);
15. Nouabale-Ndoki
II (Congo)+ Dzanga-Ndoki
II complex(Central African Republic);
16. Manovo-Gounda-St.
Floris
II** complex (Central African
Republic);
17. Salonga II;
18. Garamba
II** complex(Zaire); 19. Virunga II** (Zaire)+ Volcans I I* (Rwanda)
complex;20. Reserve de Faune Okapi(Zaire); 21. Queen Elizabeth
II* complex and Bwindi NP(Uganda);22. Mau Escarpment(not protected)(G. Davies in litt.
1993); 23. Aberdare
II (Kenya);
24. Ngorongoro
Crater VIII#(Tanzania:
IUCN 1987).

A Confirmed
record
LJ

clA

Accounts.

recordPopulation Status
Global: Category 2. Regional: Category 1. IUCN:
Insufficiently Known. While the species is tied to moist
forest habitats and is thus naturally rare, it is difficult toevaluate its conservation status due to lack of information
on its biology and ecology. The moist forests of west
Africa have been heavily degraded and remaining intact
stands are patchily distributed, while those of the Zaire
basin in Zaire, Congo, and Gabon are relatively pristineand large tracts of primary forest remain (Myers 1989,
Collins 1990, Sayer et al. 1992). However, a large portion of the latter is inland swamp forest (Sayer et al. 1992),
a habitat type in which the golden cat has not yet beenrecorded (S. Lahm in ht. 1993).
Small pieces of golden cat skin have totemic value “forwrapping things up in” (Van Mensch and Van Bree 1969,
E. Gadsby in ht. 1991). Because of taboos, people may bereluctant to discuss the animal directly (Sanderson 1940).

golden cat’s fundamental habitat-M.
Agnanga (in Zitt.
1993) reports that it is well known in northern Congo(among the most sparsely populated regions in tropical
Africa), but not in the south, where the forests are semideciduous and partially logged (Sayer et al. 1992).
Similarly, B. Hoppe-Dominik (in Zitt. 1993) describes the
species as common in the Ivory Coast’s Tai’ National Park
(rain forest), but very rare in Comoe National Park (savannah woodland).
The golden cat has been recorded at elevations up to3,600 m in Uganda (Guggisberg 1975), and in Kenya’s
Aberdare Mountains (Maberly 1966, Hardy 1979, Watson
1980). Figure 2, based on Van Mensch and Van Bree
(1969), shows the tropical rain forest of the Zaire Riverbasin as solid lines. Probable distribution elsewhere,
including patches of wet montane forest and lowland
humid forest interspersed with Savannah grasslands (former rain forest: Collins 1990), is shown as dashed lines.

11Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Cheetahby throttling it (P. Leyhausen in Ewer 1973, Kingdon
1977). Its claws remain exposed, lacking the skin sheaths
found in most other felids, and thus provide additional
traction like a sprinter’s cleats. The foot shows several
other modifications:the digital pads and also themetacarpal pad are extremely hard and pointed at the front,
possibly as an adaption to sudden braking, and the palmar
pads bear a pair of longitudinal ridges instead of the moreusual slight depressions-thefunctional equivalent of tire
treads, serving as anti-skid devices (Pocock 1916, Ewer
1973). The prominent dew claws are used as hooks to trip
up fast-running prey. The long tail helps the cheetah’s balance as it swerves during a chase. Finally, the cheetah has
enlarged bronchi, lungs, heart, and adrenals (Eaton 1974).
According to K. Sevrin (pers. comm. in Eaton 1974:24), a captive cheetah was accurately clocked at 112 kph
over a short distance. In the wild, out of 78 chases measured and timed by G. Frame (Frame and Frame 1981:181), the top speed was 87 kph. Antelopes, the main preyof cheetah, reach top speeds of 80-97 kph (Garland 1983),
so peak speeds reached at some portion of a cheetah’s
sprint probably do exceed the oft-quoted, but seldom documented, 110 kph. Cheetah sprints rarely last longer than
200-300 m, while most antelope can run much further.
Heat builds up rapidly during a sprint, and cheetahs havenot evolved the evaporative heat release mechanisms ofgazelles and goats, even though their energetic cost of running is equivalent (Taylor and Rowntree 1973, Taylor etal. 1974). Despite its refinements, the cheetah, like theother cats, is a sprinter rather than a courser.
Cheetahs are pale yellow with white underbellies, covered all over with small round black spots. They are readily distinguished from their spotted relatives by their “tearlines”-heavy
black lines extending from the inner corner of each eye to the outer corner of the mouth. Bothmelanistic and albino cheetah specimens have beenreported (Guggisberg 1975), and remarkably pale animals
have been reported from desert regions (Dragesco-Joffe
1993, P. Gros in Zitt. 1993). A more notorious single-locus
genetic mutation (Van Aarde and Van Dyk 1986) produces the blotched tabby pattern of the so-called kingcheetah (Plate 2), which was once classified as a separate
species (Pocock 1927), and was the subject of a major
investigative expedition (Bottriell 1987). This mutation
has historically been recorded only from a restricted area
in southern Africa centered on Zimbabwe (Hills andSmithers 1980), but there is a recent report of a single skinrecovered in Burkina Faso, west Africa (Frame 1992).
A greater degree of sociality has been observed among
cheetahs than for most felids, with the exception of the lion.
Male and female litter-mates tend to stay together for about
six months after independence (Caro 1994). Nearly two
decades of intensive research in the Serengeti Plains haveshown that, while females split off upon reaching sexualProtection Status
CITES Appendix II. National legislation: Fully protected
over only part of its range. Hunting prohibited: Angola,
Benin, Burkina Faso, Congo, Ghana, Ivory Coast, Kenya,
Liberia, Nigeria, Rwanda, Sierra Leone, Zaire. Hunting
regulated: Gabon, Liberia, Togo. No domestic trade controls: Congo, Sierra Leone. No legal protection: Cameroon, Central African Republic, Gambia, Guinea Bissau,
Senegal, Tanzania, Uganda. No information: Burundi,
Guinea (IUCN Environmental Law Centre 1986; M.
Agnanga, B. Hoppe-Dominik, S. Lahm in Z&t. 1993).
Principal Threats
Savannization in west Africa has probably led to population declines and fragmentation, unless there is migration
along riverine corridors. The bush meat trade, which figures largely in the region’s economy, may lead to local
depletion of small antelope prey. There appears to be little hunting of golden cats (E. Gadsby in litt. 199 1; S.
Lahm, M. Agnanga in litt. 1993).
Action Planning
Projects 20 and 21.

Cheetah, Acinonyx
Schreber, 1776

jubatus

Other Names
Guepard (French); Gepard (German); guepardo, chita(Spanish); jagluiperd (Afrikaans:South Africa); aboshamani (Amharic: Ethiopia); fahd (Arabic); bogolo
bogolo (Bournouan); marukopta (Burkina Faso); siho
(Fufulde: Cameroon); rabbi (Hausa); /uayb (Hei//kum
Bushman: Namibia); !a’o (Ju/hoan Bushman: Botswana,
Namibia); kisakasaka (Kasanga: Zaire); duma, msongo(KiSwahili);lengau, letlotse (Setswana: Botswana);
dindingwe,
ihlosi (Shona: Zimbabwe);
haramacad,daharab, horkob (Somalia); ngulule (Zulu: South Africa).
Description
and Behavior (Plate 2)
The cheetah is built for speed, with a deep chest, wasp
waist, and proportionately longer limbs than the other big
cats (Gonyea 1976). Average adult weight is 43 kg formales and 38 kg for females in the Serengeti (n=l7: Caroet al. 1987). Flexion of the elongated spine has been measured as increasing the cheetah’s stride length by 11% at
speeds of 56 kph (Hildebrand 1959, 1961). The canines
are small relative to other felids: a reduction in the size ofroots of the upper canines allows a larger nasal aperture forincreased air intake, which is critical for allowing the cheetah to recover from its sprint while it suffocates its prey12

Pat? I: Species

maturity, male litter-mates remain together in coalitions,and sometimes defend territories (Frame and Frame 1984,
Caro and Collins 1986). These coalitions, particularly trios,
may include unrelated males, with the frequency of thistype of grouping estimated at 15% in the Serengeti (Car0and Collins 1986). Males in coalitions are more likely thansolitary males to gain and maintain territories; non-territorial males live a nomadic existence and wander widely(Caro and Collins 1986, 1987a). Territorial males were
found to be in better physiological condition and appear tohave better access to females during periods of gazelle concentration (Car0 and Collins 1987b, Caro et al. 1989).
Large groups of up to 14- 19 animals (including cubs)
have been reported occasionally from parts of east andsouthern Africa where other large predators have been
eradicated (Kenya: Graham 1966, P. Gros in Zitt. 1993;
Botswana: Gros 1990; Namibia: McVittie 1979, MarkerKraus and Kraus 199 1). The advantages of grouping undersuch conditions are not clear (S. Durant in Zitt. 1993).
In east Africa, the cheetah’s main prey is theThomson’s gazelle on the plains (Serengeti: Schaller1968), and impala in the woodlands (Eaton 1974). In thearid bushland of northern Kenya, G. Adamson (inHamilton 1986a) identified lesser kudu, gerenuk, and dikdik as major prey. In southern Africa, major prey consists of springbok (northeast Botswana: Smithers 1971;
Kalahari Gemsbok National Park, South Africa: Mills
1990a; Etosha NP, Namibia: unpubl. data); greater kudu
calves and warthog (Namibian ranchland: Morsbach 1987,
L. Marker-Kraus, pers. comm.); impala (Kruger National
Park, South Africa: de Pienaar 1969, Mills and Biggs
1993); and puku (Zambia: Mitchell et al. 1965). Data arescarce for central and west Africa, but cheetahs have been
observed to take red hartebeest, oribi, and kob in ManovoGounda-St. Floris National Park in the Central African
Republic (Ruggiero 1991). Cheetahs are also known totake smaller prey, particularly
hares (Frame 1977,
Labuschagne 1979, 1981), and male coalitions often takemuch larger prey, such as wildebeest (Dorst and Dandelot
1969, Eaton 1974, McVittie 1979, Caro and Laurenson
1990, Skinner and Smithers 1990). Seasonally, a large
proportion of cheetah prey captures consist of immature
animals (McLaughlin 1970, Burney 1980). When hunting group-living
prey animals; such as Thomson’s
gazelles, they tend to select less vigilant solitary individuals (FitzGibbon 1990).
Certain aspects of cheetah behavior can be explained as
adaptations to compete with other sympatric large predators, particularly lions and hyaenas. Cheetahs are predominantly diurnal, probably because competing predators arenocturnal. It has been suggested that the cheetah’s large
litter size may be a strategy to offset high juvenile mortality caused by predators (Burney 1980, Hamilton 1986a,
Laurenson 1992, Caro 1994). Cheetahs often lose their

Accounts.

Chapter

1. Sub-SaharanAfrica,Cheetahkills to lions and hyaenas, and have only rarely been
observed to scavenge, or return to a previously abandoned
kill (Graham 1966, de Pienaar 1969, Burney 1980, Caro
1982, Stander 1990a). There is preliminary evidence thatcheetahs will remain near large kills, rather than abandon
them after satiation, on Namibian ranchlands where lions
and hyaenas have been eliminated (L. Marker-Kraus, pers.
corm-n. 1994).
Biology
Reproductive season: (W) year-round, although birth
peaks have been reported during the rainy season in the
Serengeti (November-May: Frame 1977, Laurenson et al.
1992).
Gestation: (C) 90-98 days (Marker-Kraus

1992).

Litter size: (W) 4.2 (age l-3 months) on Namibian ranchland (McVittie 1979); 3.5 (age 6-35 days; Laurenson etal. 1992) - 2.6 (age three months; Frame 1977) in the
Serengeti; (C) 3.7 (Marker and O’Brien 1989), range l-8(Green 1991).
Interbirth interval: (W) 15- 19 months (McLaughlin 1970,
Schaller 1972). Females readily go into estrus and conceive after losing a litter. Laurenson et al. (1992) foundthat the interval between the death of the previous litter andthe next successful conception was longer for young (86.3
days, n=3) than adult females (17.8 days, n=9).
Age at independence: (W) mean 18 months (Laurenson etal. 1992), range 13-20 months (Frame 1984) (sub-adults
leave mother); 17-27 months (females leave siblinggroups: Frame 1980, Laurenson et al. 1992).
Age atjirst reproduction: (W) females 24 (n=2: Schaller1972) - 36 months (n=4: Laurenson et al. 1992); males
30-36 months (Car0 1991). (C) females 2-3 years (n=lO);males l-2 years (n=8) (McKeown 1992).
Age at last reproduction: (C) females 10 years; males up to
14 years (McKeown 1992).
Sex ratio: (W) cubs: 1 male:0.95 female (n=ll7); adults
and independent sub-adults: 1 male: 1.9 females (n= 169).
This suggests differential male dispersal and mortality(Frame and Frame 1984), although males can be shyer
than females and more difficult to observe (Caro andCollins 1986).
Juvenile mortality: (W) Other large carnivores, as well asbaboons (L. Marker-Kraus in Zitt. 1993), are known to killcheetah cubs. In the Serengeti, the number of lions on thegrassy plains which constitute the Serengeti Cheetah
Project’s study area have increased tenfold since the1960s following an increase in wildebeest after rinderpest control measures. Under such circumstances, cheetah
cub mortality is very high: Laurenson (in press, pers.

13

Pat? I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Cheetahof the country, and sightings have increased further northin Awash National Park, where a cheetah was killed by atrain in 1992 (J. Hillman in Zitt. 1993).
Density and abundance vary widely according to environmental conditions, especially the occurrence of suitable prey and other large predators (Laurenson in press).
In the Serengeti Plains ecosystem, cheetahs concentrateseasonally in association with migratory movements ofThomson’s gazelle (Durant et al. 1988). Frame (1977)
found dry season concentrations of one adult per 6 km2
around woodlands/plain edge in the Serengeti. Based on
individual recognition of cheetahs from photos taken by
tourists, Bowland (1993) reported a low density for
Kruger National Park of one adult per 191 km2.
Excluding cubs under three months of age, Bumey (1980)
found that total cheetah density was twice as high in pastoral areas outside the Masai Mara National Reserve (l/29
km2) as inside the protected area (l/67 km2). On
Namibian ranchland, Morsbach (1987) reported a density of one cheetah per 50 km2.
Estimating cheetah density is complicated by their
unusual social organization.
Both solitary male andfemale adults are semi-nomadic, having large, overlapping
home ranges of the order of 800- 1,500 km2 (Frame 1980,
Morsbach 1987, Caro 1994). Coalitions of males, on theother hand, have been found (in the Serengeti) to defend
small territories of the order of 12-36 km2, but up to 150
km2 (Bertram 1978, Frame 1980, Caro and Collins 1986).
These territories periodically hold high numbers ofThomson’s gazelle, the favored prey of female cheetahs,
and females were often observed in the males’ territories(Car0 and Collins 1987b).

comm. 1993) found that 73% of cub deaths were due topredation (mainly lion), and that a total of 95% of 125 cubs
failed to survive to independence.
Longevity: (W) 12-14 years (Frame and Frame 1980).
However, Laurenson (in press) estimates the mean life
expectancy of females reaching three years of age in the
Serengeti at only an additional 3.9 years. Territorial males
probably live longer, on average, than single males (Car0and Collins 1986, Caro et al. 1989). (C) average 10.5 andup to 21 years (L. Marker-Kraus in Zitt. 1993).
Habitat and Distribution
Cheetahs are distributed primarily throughout the drierparts of sub-Saharan Africa (Fig. 3). They are not generally associated with forest habitats: they occur only thinly
in the more humid zones of miombo woodland that covermuch of central southern Africa, and are absent from theSudano-Guinean forest Savannah belt of west Africa(Myers 1975). However, although cheetahs are most frequently observed on open grassy plains (e.g. Schaller
1972, Mills and Biggs 1993), they also make extensive useof bush, scrub, and open woodlands (Myers 1975,
Hamilton 1986a, Morsbach 1987). Observations by Eaton( 1974) suggest that cheetahs expend more energy hunting
in open country than in cover. A mosaic of woodland andgrassland is probably preferred. They range up to 1,500
m in the mountains of Ethiopia (Yalden et al. 1980).
Cheetahs are well-adapted to living in arid environments. They are not obligate drinkers and, in the Kalahari
desert, have been estimated to travel an average of 82 km
between drinks of water. They were observed to satisfy
their moisture requirements by drinking the blood or urineof their prey, or by eating tsama melons (Labuschagne
1979, 1981).

Protection Status
CITES Appendix I. An Appendix I quota system wasestablished under CITES in 1992 for live animals and trophies, with annual quotas allocated as follows:
150(Namibia), 50 (Zimbabwe), 5 (Botswana). National legislation: fully protected over most of its range. Hunting
prohibited: Angola, Benin, Botswana, Burkina Faso,
Cameroon, Central African Republic, Ethiopia, Ghana,
Kenya, Malawi, Mali, Mauritania, Mozambique, Namibia,
Niger, Nigeria, Rwanda, Senegal, South Africa, Sudan,
Tanzania, Togo, Uganda, Zaire. Trophy hunting permitted: Namibia, Zambia, Zimbabwe. No information: Chad,
Sudan (IUCN Environmental Law Centre, 1986; Kraus
and Marker-Kraus 199 1).

Population Status
Global: Category 3(A). Regional: Category 2(A). IUCN:
Vulnerable. The total number of cheetahs in sub-SaharanAfrica has been variously estimated at 15,000 (Myers
1975), 25,000 (Frame 1984), and 9,000-12,000 (Kraus
and Marker-Kraus
1991), and a wide-ranging survey isin progress to develop a better grasp of the cheetah’s current status (P. Gros, in prep.). The two largest metapopulations of cheetah are now believed to occur in east
Africa (Kenya and Tanzania) and southern Africa(Namibia, Botswana, Zimbabwe and Zambia) (Kraus andMarker-Kraus
1991, Gros 1990 and in Zitt. 1991). The
cheetah appears to be most rare in the Sahelian and
Sudanian semi-arid zones of west Africa-althoughoriginally optimal habitat, much of it is now very degradedunder human population pressures (T. Anada in Zitt.
1993). Cheetah populations are still believed to behealthy in Ethiopia, with their stronghold across the south

Principal

Threats

Genetic homogeneity: Genetic research has demonstrated
that both captive and free-ranging cheetahs exhibit a very
high level of homogeneity in coding DNA, on a par withinbred strains of laboratory mice (O’Brien et al. 1983,

14

Part I: Species

1985, 1986, 1987a). The cheetah appears to have suffereda series of severe population bottlenecks in its history, withthe first and most significant occurring possibly during
the late Pleistocene extinctions, around 10,000 years ago

Accounts.

Chapter

1. Sub-SaharanAfrica,Cheetah(Menotti-Raymond and O’Brien 1993). The factors which
would have led to these ancient population bottlenecks
are not clear, but both their causes and consequences could
be of significance to cheetah conservation today.

Figure 3. Distributionof the cheetah(A. jUx&~s)in sub-SaharanAfrica.
1. Boucle du Baoule II complex; 2. Adrar des Iforas Mts Reserve (proposed:Mali); 3. “W” II* complex(Burkina
Faso, Benin and Niger); 4. Air and Ten&-e VIII; 5. Termit Massif (not protected)(Niger);
6. Benoue II* (Cameroon);
7. Ouadi Rime-Ouadi
Achim Fauna1 Reserve;
8. Zakouma
II complex (Chad);
9. Manovo-Gounda-St.
Floris II** complex (Central African Republic);
IO. Ouandjia-Vakaga
IV (Central
African Republic) + Radom II* (Sudan);
11. Dinder II* complex;
12. Boma II (Sudan);
13. Mago + Omo II
complex;14. Gambella
V; 15. Yangudi Rassa II; 16. Awash II (Ethiopia);
17. Kidepo Valley II complex(Uganda);18. Kora II complex;
19. Tsavo II complex (Kenya);
20. Maasai Mara II (Kenya) + Serengeti
II# (Tanzania)
complex;21. Ruaha II complex;
22. Selous IV** complex (Tanzania);
23. Kasungu II(Malawi);24. Kundelungu
II complex (Zaire); 25. South Luangwa
II complex;
26. Sioma Ngwezi II;
27. Kafue II complex (Zambia);
28. Mana Pools II** complex; 29. Hwange II complex;
30. Gonarezhou
II (Zimbabwe);
31. Kruger II complex;
32. HIuhIuwe/Umfolozi
IV and other Natal reserves;
33. Pilanesberg II (reintroduced)(South Africa); 34. Kalahari Gemsbok
II (South Africa) + Gemsbok
II (Botswana)complex;35. Central Kgalagadi
II (Botswana);
36. Namib-Naukluft
II; 37. Etosha II (Namibia);
38. lona
VI; 39. Bikuar II; 40. Luando IV; 41. Kisama V (Angola);
42. The cheetah is consideredvery rare if not
extirpatedin Nigeria, but there was a recent sighting from Falgore (Kogin Kano) IV (F. Hurst in Mt. 1994);
43. Gadabedji
IV (Niger).

15

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Cheetah

It has been argued that lack of genetic diversity may
render the cheetah an exceptionally vulnerable species(O’Brien et al. 1983). Genetic variation is thought to be
essential to the long-term adaptability and persistence of
populations by providing sufficient genetic options on
which natural selection can operate in response to environmental change. The evidence for cheetahs being compromised by their genes arises mainly from captivity,
where epidemics of infectious disease have occurred withhigh mortality (O’Brien et al. 1985, Evermann et al. 1988).
Increased susceptibility to disease has been linked togenetic monomorphism (O’Brien and Evermann 1988).
Zoos have had great difficulty in breeding cheetahs.
Captive female cheetahs conceive infrequently, and when
they do, cub mortality is relatively high (28-36%) (Marker
and O’Brien 1989; Marker-Kraus
and Grisham 1993),
although these rates are similar to those of other felid andcarnivore species kept in captivity (Loudon 1985).
Finally, both wild and captive male cheetahs have highlevels of abnormal sperm (71-76%: Wildt et al. 1987a),and success with in vitro fertilization using cheetah sperm
is relatively low compared to other felid species(Donoghue et al. 1992).
However, there is no evidence that reproduction is compromised in the wild (Caro and Laurenson 1994). To alarge extent, the cheetah’s poor reproductive performancein captivity is linked to institutional management practices.
First, some zoos have had high success in breeding cheetahs (Van Dyk 199 1, Lindburg et al. 1993). Factors whichappear to facilitate breeding include large enclosures withlong views, constant separation and reintroduction ofmales and females, and provision of secluded nest boxes
for mothers with young (Lee 1992, Laurenson 1993).
Second, vulnerability to disease increases in captive situations, and no epidemics have been reported from wild populations, although cheetahs in some parks h.ave beenreported to suffer a relatively high incidence of mange(Caro et al. 1987, Bowland 1993, R. Kock in litt. 1993).
Finally, some captive males are very fertile and others
essentially infertile, despite having similar levels of poor
quality sperrn (Donoghue et al. 1992, Lindburg et al. 1993,
Wildt et al. 1993a).
The cheetah’s genetic monomorphism is a fascinating
aspect of its bi ology, and poten tially of importance to its
conser ,vation, but implications for management of wild
POPUlations are not yet evident.

ards, and hyaenas-limit
cheetah abundance is by killing
cheetah cubs (Laurenson in press), but these species, aswell as (sometimes) jackals, baboons, and vultures, also
drive adult cheetahs off their kills. The cheetah’s relatively
large litter size may be a strategy to offset high juvenile
mortality (Burney 1980, Hamilton 1986a, Laurenson
1992, Caro 1994). Where other large carnivores havelargely been eliminated, such as ranchland in Namibia,
farmland and pastoral land in Kenya, and in parts ofSomalia, cheetahs appear to flourish at higher densities(McVittie 1979, Burney 1980, Hamilton 1986a, Morsbach
1987, A. Simonetta in Zitt. 1993). A strategy of relying
solely upon the limited system of protected areas within
the cheetah’s range may not be sufficient to ensure theconservation of viable sub-populations.
Livestock Predation: The survival of the cheetah outside
protected areas is affected by conflicts with people overpredation on livestock. Cheetahs are reported to prey on
young camels and goats in the Air and Termit regions ofNiger (T. Anada in Zitt. 1993). In Namibia, the cheetah isviewed as the most important predator of livestock on bothcommercial and communal farms: annual losses for thesefarms have been reported at lo- 15% for small stock (sheep
and goats) and 3-5% for cattle calves up to eight months ofage (Morsbach 1984-1986). Inevitably, stock losses topredators are greater where the natural prey base has been
eliminated or reduced: on a 200 km2 ranch in Kenya,
where about 9,500 head of livestock graze alongside a stilllargely intact wild ungulate assemblage, depredation bycheetahs is minimal, accounting for only 11 sheep a year
(Mizutani 1993).
Although farmers’ estimates of stock losses to cheetah
may be inflated, either intentionally or otherwise, the fact
remains that the species is widely considered a threat topeople’s livelihood, and governments have little hope of
preventing the destruction of cheetah on private lands if thatis what the owners wish to do. Hamilton (1986a) points outthat the cheetah may be more resilient to eradication onranchland than other large carnivores-which
will, forexample, take poisoned bait-but the cheetah’s decline on
Namibian ranchland during the 1980s is certainly attributable to persecution (Morsbach 1987). Namibia, South
Africa, and Zimbabwe are now pursuing a strategy of permitting trophy hunting of cheetahs on private land, with thegoal of encouraging landowners to accept and profit fromcheetahs on their land. In addition, the Cheetah Conservation Fund of Namibia is working to educate farmers about
appropriate management steps that can be taken to minimize stock losses (see Part II Chapter 2).

Vulnerability in Protected Areas: Many observers havecommented on the cheetah’s vulnerability to interspecificcompetion with other large carnivores, and this is now theprimary focus of the long-term cheetah study in the
Serengeti (S. Durant, pers. comm.1993). The chief mechanism by which more powerful carnivores-lions,leop-

Action Planning
Projects 22-27.

16

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Africanlion

second when extensive culling of lions in the Kruger
National Park opened up large vacancies for immigration(Smuts 1978a). Pride sizes (measured by the number ofadult females) are smallest in very arid environments(mean 2.2 in South Africa’s Kalahari Gemsbok National
Park: Eloff 1973a) and otherwise average between fourand six (Schaller 1972, Smuts 1976, Hanby and Bygott
1979, Ruggiero 199 1, Stander 199 1). Pride size is positively correlated with lean season prey abundance, and in
the Ngorongoro Crater, where prey is abundant yearround, groups of up to 20 adult females have been
observed (van Orsdol et al. 1985).
Prides are “fission-fusion”
social units: membership isstable (for example, three prides in the Serengeti haveoccupied the same ranges for more than 20 years), but the
pride members are often scattered in small sub-groups
throughout the pride’s range, and each individual spends
a considerable amount of time alone (Schaller 1972,
Bertram 1978, Pusey and Packer 1987). Females demonstrate several cooperative behaviors unique among the
felids. Pride members often give birth in synchrony, andthe young are reared communally, with cubs suckling
freely from lactating females (Schaller 1972, Rudnai 1974,
Bertram 1975b). Groups of females do most of the hunting, and males, for the short time that they are living
together with females, concentrate their energy on defending their tenure (see below). Stander (1992a) found thatmales in Namibia’s Etosha National Park failed to participate in hunts in 96% of 461 opportunities.
In general, prides often divide into smaller sub-groups
when foraging (range l-7: Stander 1992a, Scheel 1993).
Stander (1992b) found a complex division of labor among
hunting lionesses, with individuals repeatedly playing thesame role of either “center” or “wing.”
Centers, which
tended to be larger and heavier lionesses, generallyambushed and captured prey chased by the wings.
However, lionesses were flexible and would switch roles,
depending on group composition and positioning.
D.
Joubert (in Zitt. 1993) suggests that lionesses also switch
roles according to prey type: in Botswana’s
SavutiNational Park, he has observed, “with some consistency,”
the same lioness take the lead in warthog hunts, while
playing a passive role in buffalo hunts.
A single male or coalition of males (up to seven) holds
tenure over one or more prides, and effectively excludes
strange males from siring cubs with pride females (Packer
et al. 199 1a). Competition among males for pride tenure isintense, and average tenure is only two (Packer et al. 1988)
to three years (Stander 1991). Males will only seek tenure
over or breed with related pride females under unusual
circumstances (e.g.when the population is small and there
are barriers to dispersal: Pusey and Packer 1987, Packer
et al. 1991a,b). Males are also highly social: coalitions in
the pre- and post-tenure periods hunt and scavenge coop-

African lion, Panthera leo(Linnaeus, 1758)
Other Names
Lion d’Afriqued’Afiique (French); Lowe (German); lean (Spanish);ambessa (Amharic: Ethiopia); nkharam (Chichewa:
Malawi); xamm (Damara: Namibia); zaki (Hausa); odum,
aja (Ibo, Yoruba: Nigeria); n!hai (Juhoan(Ju/hoan Bushman:
Botswana, Namibia); ngatia, muruthi (Kikuyu: Kenya);ngouambulu (Lingala: west Africa); labwor (Luo: Kenya,
Uganda); olugatany (Maasai, Samburu: Kenya, Tanzania);
leao (Portugese); tau (Setswana: Botswana);
simba(KiSwahili); aar, baranbarqo, libaax, gool, davar (Somalia).
Description
and Behavior (Plate 1)
Smuts (1976) reports the following weight series for lions
in South Africa’s Kruger National Park: adult males (>4years) 181 kg (n=14) and females 126 kg (n=25); sub-adultmales (2-4 years) 146 kg (n=25) and females 103 kg(n= 11); large male cubs (l-2 years) 77 kg and females 60
kg. The largest adult male weighed 225 kg, and the largest
female 152 kg (Smuts 1982). A male shot near Mount
Kenya in 1993 weighed 272 kg (R. Kock in Zitt. 1993).
The record total lengths (including the tail) for male lions
are around 3.3 meters (Guggisberg 196 1). Lions have uniformly tawny coats. While the color may vary locallyfrom pale to dark, leucism (unusual white pelage but withpigmented eyes and skin, as opposed to true albinism
which is a complete lack of pigmentation) has beenreported only from the vicinity of Kruger National Parkand the Umfolozi Game Reserve in South Africa(McBride 1977, Smuts 1982), and a black form has neverbeen observed (Guggisberg 1975). Lions are the only catswith tufted tails and manes (males only). The maneappears to serve several functions: increased protection
during intraspecific fighting; a signpost of gender distinguishable at distance (possibly linked to the lion’s historiccolonization of open plains); and an indicator of individual
fitness (Schaller 1972, Kingdon 1977). The males ofmany polygynous species tend to develop conspicuous display features. The fact that only the lion, out of all cats,has done so suggests that the mane is closely linked to thelion’s distinctive social system. Mane development isstrongly influenced by testosterone (Schaller 1972).
The core unit of the lion’s matrilocal society is the
pride, which consists of a group of related females (nonedominant) and their cubs (Schaller 1972, Bertram 1975a,
Packer et al. 199 1a). There are only two recorded casesof unrelated females forming a pride, and both cases
involved prides giving up their original natal ranges: thefirst when prolonged severe drought in Botswana’s Central
Kalahari Game Reserve rendered the females’ original
ranges uninhabitable (Owens and Owens 1984), and the17Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Africanlion

eratively, and larger coalitions of 4-6 males can maintaintenure more than twice as long as l-2 males (> 47 months)(Bygott et al. 1979).
Despite maternal defense, infanticide is common whenmales take over a new pride: most females with dependent offspring lose their cubs within a month of a takeover,and those that are pregnant lose their cubs shortly after giving birth. In this way, males assure paternity during theirshort reproductive lifetime, which is generally only as longas their period of pride tenure. In response, females show
a burst of heightened sexual activity for about threemonths following a takeover, attracting other males andencouraging competition that ensures that the fittest (often
largest) coalition is able to gain tenure. They remain infertile (anovulatory: Smuts et al. 1978) during this “testing”
period, and only afterwards, when tenure has stabilized,
tend to breed in synchrony (Packer and Pusey 1983).
Litters born synchronously have a higher survival rate
(probably due to maximal maternal care [Bertram 1975b]),and tend to show a sex ratio biased toward males. Thismay be because groups of related males reproduce more
successfully (Pusey and Packer 1987).
Coalitions of >4 males are always related (being born inthe same pride, but not necessarily of the same mother),while pairs frequently consist of unrelated males (and lessfrequently, a related pair teams with an unrelated male toform a trio) (Packer et al. 1991 a). Reproductive successincreases with coalition size (Bygott et al. 1979, Packer etal. 1988). Although at least one member of male coalitions larger than two fails to breed successfully (Packer etal. 199 1a), through kin selection (Bertram 1976) nonbreeding helpers which are related still ensure that some
portion of their genes are passed down.
The question of why sociality evolved to such a highdegree in lions has been the subject of considerable debate.
There were probably several contributory causes, which
occurred many generations ago. Data from present-day
studies cannot refute any of them, but can shed some lighton how and in what circumstances they might work (B.
Bertram in ht. 1993). Evidence suggests that coordinatedgroup hunts are more successful at capturing (Packer andRuttan 1988, Stander 1992a,b) and killing (Packer 1986)very large prey (see below for discussion of major preyspecies). Stander and Albon (199’2) found that hunting
success, even for smaller antelope prey, increased linearlywith foraging group size in the semi-arid open plains ofEtosha National Park. However, what would seem to be
the most obvious explanation-increased
hunting successyields more food- becomes less so on closer examination.
Even on large carcasses, it appears that the presence ofnumerous non-hunting “cheaters” (Packer and Ruttan
1988) within the pride can reduce per capita food intake
to the point where cooperative hunting does not appear tobe economic for the hunters. The highest rate of foodintake per hunt appears to be gained by solitary females(Packer 1986). Packer (1986), based on the theory of kin
selection, argued that lions became social because it is evolutionarily more advantageous to share kills with scavenging relatives than to yield to strange lions or other large
predators. Other benefits of sociality have also been
pointed out: defense of young, maintenance of long-term
territories (Packer et al. [ 1990]), insurance against individual injury or incapacity, and minimization of chances ofgetting no food at all (B. Bertram in ht. 1993).
Major large ungulate prey species recorded in east, central, and southern Africa include buffalo, zebra, wildebeest, roan, sable, springbok, gemsbok, kob, impala,warthog, waterbuck, and hartebeest (Mitchell et al. 1965,
Makacha and Schaller 1969, de Pienaar 1969, Schaller
1972, Eloff 1973a, Rodgers 1974, Rudnai 1974, Bertram
1978, Berry 1981, van Orsdol 1982, 1984, Smuts 1982,
McBride 1984, Mills 1984, Fagotto 1985, Prins and Iason
1989, Ruggiero 1991, Stander 1992a, Scheel 1993, Viljoen
1993). While medium to large-sized ungulates make up
the bulk of their diet, lions, like leopards, are generalist
hunters, and will take a wide range of prey, from smallrodents (Eloff 1973a) to young rhinos, hippos, and elephants (McBride 1990, Ruggiero 1991, Viljoen 1993; H.
Dublin, H. Jachmann in ht. 1993). Individual differencesin prey selection and killing techniques are often discernible for different prides in the same area (Rudnai 1973,
van Orsdol 1984, McBride 1990, Mills and Biggs 1993),
indicating a strong role for learning in the lion’s hunting
behavior. For example, a pride of lions which occasionallyforaged along Namibia’s Skeleton Coast desert learned
how to prey and scavenge upon Cape fur seals (Bridgeford
1985, Berry 1991a). (The entire pride was eliminated in1991 by cattle herdsmen [Berry 1991b., L. Scheepers,
pers. comm. 19931). Lions (especially males) frequently
scavenge (>40% of food items in the Serengeti: Packer etal. 1990), although this behavior is less common in arid
environments, where prey occurs at lower density (4.6[Mills 1990 - 6% [Stander 1992a] of food items).
Lions usually (but not always) hunt at night (Schaller
1972, van Orsdol 1982, Mills and Shenk 1992, Stander
1992a). In Botswana’s Savuti National Park, D. Joubert(in Zitt. 1993) reported a higher success rate when lions
hunted on moonless nights. Their distinctive roar, which
in optimal conditions can be heard up to five km away
(Guggisberg 1975), appears to serve to demarcate territories (Schaller 1972), much as scat deposits do for the otherlarge cats. Stander and Stander (1987) found it possible
to distinguish between not only the roars of males andfemales, but also of individual males.
Outside protected areas, where lions are heavily persecuted and the wild ungulate prey base is reduced, groupsizes are reported to be much smaller (l-2: Thomas 1990,
F. Hurst in ht. 1991), and they are seldom heard to roar18

Part I: Species(Thomas 1990, C. Stuart in Zitt. 1991). It is not clear
whether the lion’s social system “breaks down” under suchconditions of low prey and low lion density. Small foraging group size may be more efficent for stock-raidinglarger groups would be more conspicuous and vulnerable
to rancher retaliation (H. Dublin, C. Packer in Z&t. 1993).

Accounts.

Chapter

1. Sub-SaharanAfrica,Africanlion

able to eat well from group-shared carcasses (Schaller
1972, van Orsdol et al. 1985). Infanticide is also an important factor (Packer and Pusey 1983). Van Orsdol et al.
(1985) reviewed cub mortality (< 12 months) across a
range of habitats: rates ranged from 14-73%.
Sex ratio: (W) Prenatal: 1 male:0.9 female (n=39); adult(5+ years): 1 male:2.1 females (n=373 lions, Kruger NP:
Smuts 1978b). Adult sex ratios are typically heavily
weighted in favor of females. The skew does not appear to
be related to food supply or density, but rather to differential rates of maturation, mortality, and emigration between
the sexes (van Orsdol et al. 1985).

Biology
Reproductive season: (W) Largely aseasonal (Bertram
1975b), but weak (February-April: Smuts et al. 1978) andstrong (March-July:
Packer et al. 1990) birth peaks
recorded in Kruger and Serengeti National Parks.
Estrus: (W) 4 days.

Age at last reproduction: (W) female reproductive performance starts to decline at 11 years and virtually ceases at
15 (Packer et al. 1988); 16 year-old males can still produce
viable sperm (Smuts et al. 1978), but reproduction probably completely ceases after pride tenure is lost (8- 10 years:
Packer et al. 1988).

Interestrus interval: (W) 16 days (Packer and Pusey 1982).
Gestation.- (C) mean 110 days (range 100-l 14; n=51)(Cooper 1942).
Litter size: (W) from field counts of small cubs < 1 yearof age, average 2.5 (n=59, Serengeti NP: Bertram 1975b) 3.02 (n=47, Kruger NP: Smuts et al. 1978); range 1-6, but
98% of litters are l-4 (n=274, Serengeti NP: Packer andPusey 1987).

Longevity: (W) males generally 12 (Hanby and Bygott1991), and up to 16 years (Smuts et al. 1978), females generally 15- 16 (Hanby and Bygott 199 I), and up to 18 years(Bertram 1975a); (C) average 13 years, but up to 25-30(Guggisberg 1975).

Interbirth interval: (W) mean 20 months (range 1 l-25;n=38) if previous litter survives to maturity (12 months);
4-6 months if previous litter lost (Pusey and Packer 1987).

Habitat and Distribution
Optimal habitat appears to be open woodlands and thickbush, scrub, and grass complexes where sufficient coveris provided for hunting and denning. The lion has a broad
habitat tolerance, absent only from tropical rain forest andthe interior of the Sahara desert. Although lions drink regularly when water is available, they are capable of obtaining their moisture requirements from prey and even plants(such as the tsama melon in the Kalahari desert), and thus
can survive in very arid environments (Eloff 1973b).
They may range quite high into the mountains of east
Africa, up to 3,600 m on Kenya’s Mt. Elgon (Guggisberg
1961), and 4,240 m in Ethiopia’s Bale Mountains (Yalden
et al. 1980).
The lion formerly ranged from northern Africa throughsouthwest Asia (where it disappeared from most countries
within the last 150 years), west into Europe, where it
apparently became extinct almost 2,000 years ago, and east
into India (where a relict population survives today in the
Gir Forest: see species account in North Africa andSouthwest Asia) (Guggisberg 1961). Lions survived in thedesert on the edge of Niger’s Air Mountains up to about 60
years ago (Rosevear 1974).

Age at dispersal: (W) Males generally leave their natal
pride at between 2-4 years (Schaller 1972, Bertram 1975a,
Pusey and Packer 1987), but young males may be forcedout much earlier by a pride takeover, e.g. 13-20 months(Hanby and Bygott 1987). Most young females are incorporated into their natal prides, but about 33% disperse toform new prides in the Serengeti (Pusey and Packer 1987).
The percentage of dispersers may be higher elsewhere (D.
Joubert in Zitt. 1993). Median age at dispersal for femalesis 2.5 years (75% of dispersers between 1.5-3.75 years ofage: C. Packer in litt. 1993).
Age atfirst reproduction: (W) While the onset of spermatogenesis begins at 30 months in males (Smuts et al.
1978), and females may begin mating at 24 months, successful first reproduction generally happens only when
pride membership is established. In the Serengeti, females
which remained within their natal pride first gave birth atfive years (n=22). Females which dispersed from theirnatal pride first successfully raised litters at an average ageof 8 years (n=8 emigrant cohorts); earlier litters generally
did not survive. Males generally establish pride tenure at 44.5 years, with larger coalitions (4+) establishing residence
earlier (Pusey and Packer 1987).

Population Status
Global: Category 3(A). Regional: Category 2(A). IUCN:
not listed. There are no sound estimates of the total number of lions in Africa: guesstimates range from 30,000 to

Cub mortality: (W) Mortality of cubs is rather high inlions, and is linked chiefly to periods of prey scarcity,when kills may be more infrequent and cubs may not be

19

Par? I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Africanlion

100,000 (Stuart 1991, P. Jackson, pers. corm-n.). East andsouthern Africa are home to the majority of the continent’s
lions; in west Africa, numbers have greatly declined.
Throughout most of Africa, lions are becoming increas-

ingly rare outside protected areas (Fig. 4).
The countries in which lions are still relatively widespread are Botswana, Central African Republic, Ethiopia,
Kenya, Tanzania, Zaire, and Zambia. Status in Angola,

Species range: reduced abundance,
populations
highly localized

Figure 4. Distributionof the lion (I? lea) in sub-SaharanAfrica.
1. Niokola-Koba
II# (Senegal);
2. Boucle de Baoule II complex(Mali); 3. Comoe II# (Ivory Coast);
4. Mole II (Ghana);
5. Kabore-Tambi
II (Burkina
Faso); 6. “W” II* complex (Burkina
Faso, Benin and
Niger); 7. Kainji Lake II; 8. Kwiambana
VIII complex; 9. Lame/Burrs
IV complex;
IO. Yankari II;
11. Chingurmi/Duguma
Game Reserve(Nigeria);
12. Waza II*; 13. Benoue II* (Cameroon);
14. Zakouma II complex (Chad); 15. Nana-BaryaIV; 16. Bamingui-Bangoran
II* complex;
17. Manovo-GoundaSt. Floris II** complex (Central African Republic);
18. Simien Mts. II**; 19. Gambella
V; 20. Bale Mts. II;
21. Mago + Omo II complex (Ethiopia);
22. N and S Karamoja
VI complex (Uganda);
23. Sibiloi II;
24. Tsavo II complex (Kenya);
25. Maasai Mara II (Kenya) + Serengeti
II# (Tanzania)
complex;
26. Selous IV** complex;
27. Moyowosi
IV (Tanzania);
28. Bili-Uere VI; 29. Virunga II** complex;
30. Upemba
II + Kundelungu
II complex (Zaire); 31. Odzala II* complex (Congo);
32. Kisama II;
33. Luando IV; 34. lona II; 35. Bikuar II (Angola); 36. Etosha II; 37. Kaudom VIII; 38. West Caprivi IV(Namibia);
39. Kafue II complex; 40. Mweru-Wantipa
II complex; 41. N and S Luangwa
II complex(Zambia);
42. Liwonde II (Malawi);
43. Rovuma (Niassa) Game Reserve;
44. Banhine NP (Mozambique); 45. Kruger II complex;
46. Hluhluwe-Umfolozi
IV complex (South Africa); 47. Gemsbok
II(Botswana)+ Kalahari Gemsbok
II (South Africa) complex; 48. Central Kgalagadi
II complex;
49. Chobe II (Botswana).

20Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,ServalZambia, Zimbabwe. No legal protection: Burundi, Guinea
Bissau, Lesotho, Namibia, Swaziland, South Africa. No
information: Burundi, Chad, Djibouti, Guinea (IUCN
Environmental Law Centre 1986).

Mozambique, Sudan, and Somalia is difficult to determine because of these countries’ long history of civil
unrest; in Angola lions are believed to be widespread but
rare (Anstey 1992), and in Somalia they are patchily distributed, and largely restricted to the south (Fagotto 1985,
A. Simonetta in Zitt. 1992).
Populations are well-defined, but isolated and centeredon protected areas in the following southern African countries: Namibia (Etosha NP 300; northeastern region 130200; Caprivi Strip 40-60; northwestern region 35-40 [H.
Berry, P. Stander in Zitt. 19911) and Zimbabwe (Hwange
National Park complex 500; Gonarezhou National Park
complex 200; Zambezi Valley and Sebungwe complexes
300 [Stuart and Wilson 19881).
Lions are more sparsely distributed in Benin, Burkina
Faso, northern Cameroon, southern Chad, southern
Congo, northern Ivory Coast, northern Ghana, northern
Guinea, eastern Guinea Bissau, southern Mali, northern
Nigeria, and Uganda. Populations are essentially restricted
to protected areas only in Burundi, Malawi, Niger,
Rwanda, Senegal, and South Africa. Lions are believed
to be extinct or practically so in Djibouti, Gabon(Franceville area), Lesotho, Mauritania, Swaziland, andTogo (Limoges 1989, Stuart 1991; E. Abe, M. Agnanga,
T. Anada, A. Blom, P. Chardonnet in Zitt. 1993).
Reported lion densities (measured according to numbers of adults and sub-adults per 100 km2) range from 0.17
in the Savuti region of Botswana’s Chobe National Park
(Viljoen 1993) to 1S-2 (Kalahari Gemsbok NP: Mills etal. 1978; Etosha NP: Stander 1991) to 3-10 and up to 18
in east and southern African protected areas (Makacha and
Schaller 1969, Schaller 1972, Rudnai 1973, Rodgers 1974,
Smuts 1976, van Orsdol et al. 1985, H. Jachmann in ht.
1993). The highest known density is in Kenya’s Maasai
Mara National Reserve, the northern extension of the
Serengeti plains ecosystem, estimated at 301100 km2 (H.
Dublin in ht. 1993). Density is closely linked to seasonal
prey availability (van Orsdol et al. 1985). Average pridehome range sizes vary from 26 to 226 km2 (van Orsdol etal. 1985, Viljoen 1993), and can be considerably largerStander (1991) reported that one pride in Etosha NP had ahome range of 2,075 km2.

Principal Threats
Lions are generally considered serious problem animals
whose existence is at odds with human settlement and cattle culture. Their scavenging behavior makes them particularly vulnerable to poisoned carcasses put out toeliminate predators (E. Abe, T. Anada, P. Chardonnet, A.
Simonetta in ht. 1993). Where the wild ungulate preybase is migratory, stock-raiding has been reported toincrease during the lean season (H. Dublin in Zitt. 1993).
Problems of managing big cats in the vicinity of humansettlement are discussed in Part II, Chapter 2.
Action Planning
Projects 28-32.

Serval, Leptailurus(Schreber, 1776)

servalOther Names
Serval, chat-tigre, lynx tachete (French); Servalkatze(German); serval (Spanish); tierboskat (Afrikaans: South
Africa); aner (Amharic: Ethiopia); amich boudrar, ouchiak
zilagla (Berber: Kabylia, Algeria); njuzi (Chichewa:
Malawi); onca de baga baga (Creole: Guinea-Bissau);!‘hbm!a (Ju/hoan Bushman: Botswana, Namibia); gato
serval, gato lagar (Portugese); tadi (Setswana: Botswana);
muq shabeel, dumad xabashi, shabeel adari, shabeel yer(Somalia); mondo (KiSwahili);ingwenkala,indlozi(Xhosa, Zulu: South Africa).
Description
and Behavior (Plate 2)
The serval is well-adapted to hunting small prey in longgrass: its legs are slim and relatively long, and shoulder
height is about 0.6 m. Its neck is also elongated, its head issmall and delicate, and its ears are tall. The auditory bullae are correspondingly well-developed, making up about
22% of skull length (Skinner and Smithers 1990). Males
weigh 9- 18 kg (averaging 1l- 13 kg), and females 9- 13 kg,(averaging 9.7-l 1 kg: Smithers 197 1, Kingdon 1977,
Smithers 1978). Coat color is pale yellow, and is marked
with solid black spots along the sides and bars on the neckand shoulders.
Although 17 subspecies are listed by Allen (1939), theirvalidity is doubtful (see Appendix 1). Smithers (1978)
examined specimens from one locality in southern Africa
and found external characters among them which had beenused to designate six different subspecies within the sub-Protection Status
CITES Appendix II. National legislation: hunting
restricted to “problem” animals over much of its range;some trophy hunting. Hunting prohibited: Angola, Cameroon, Congo, Gabon, Ghana, Malawi, Mauritania, Niger,
Nigeria, Rwanda. Hunting regulated or restricted to “problem/dangerous” animals: Benin, Botswana, Burkina Faso,
Central African Republic, Ethiopia, Ivory Coast, Kenya,
Mali, Mozambique, Senegal, Somalia, Sudan, Tanzania,
Togo, Uganda, Zaire, Zambia, Zimbabwe. Trophy hunting
permitted: Botswana, Namibia, South Africa, Tanzania,

21

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Servalble-although
the study animals were habituated, they
were not radio-collared), Geertsema (1985) found that
adult males, adult females and sub-adults spend about 25%of each 24-hour period travelling and hunting. On average, Ngorongoro servals killed about 16 times within thisperiod. Independent sub-adults killed more frequently
than adults, but took smaller prey with a lower energetic
return. From nearly 2,000 observations of pounces,
Geertsema (1985) found serval hunting success to average 49%, with no significant difference between day andmoonlit night. After giving birth to kittens, one female
increased her success to 62% from 48%.

region. Servals from west Africa most frequently show apattern mutation of small speckled spots-theseso-calledservalines were considered a separate species (FeZisbrachyura Wagner, 1841) until Pocock (19 17a) demonstrated that the speckled form was a serval morph. Black
servals have been widely recorded (Shortridge 1934, York
1973, Guggisberg 1975). The holotype of L. sewal was
taken near the Cape of Good Hope, but the serval now
appears to have been extirpated from the entire southerncoastal belt of South Africa and most of Cape province(Skead 1980, Stuart 1985)-although M. Bowland (in litt.
1993) notes an unconfirmed report from a farmer at
George, midway between Cape Town and Port Elizabeth.
Small mammals, especially rodents, are the serval’s
main prey. Larger rodents are preferred, particularly vlei
(swamp) rats (Smithers and Wilson 1979, Geertsema
1985, Bowland 1990), and Nile rats (Geertsema 1976,1985). Smaller mice are of secondary importance(Smithers and Wilson 1979, Geertsema 1985, Bowland
1990). Up to 12 mice were found in one serval stomach
from Zimbabwe (Smithers 1978). Birds, reptiles, fish, andinsects are also taken, although infrequently when rodents
are abundant (Geertsema
1985, Bowland
1990).
Geertsema (1985) observed one young male serval, on a
moonlit night, rush into open water to seize one of a groupof feeding flamingos. Geertsema (1985) also found frogs
to be a particularly favorite prey item, with remains occurring in 77% of 56 scats. She saw another young male eat
at least 28 frogs in one three-hour period. Servals do not
generally take larger prey as does the caracal. Single animals have only rarely been observed to kill duikers andfawns of the smaller antelope species (Rahm 1966, dePienaar 1969, York 1973). The detailed studies byGeertsema ( 1985: Ngorongoro
Conservation
Area,
Tanzania) and Bowland (1990: Natal province farmland,South Africa) did not record any instances of servals taking mammalian prey 1;irger than rodents.
The serval locates prey in tall grass or reeds primarily
by hearing. It makes a characteristic high leap as it
pounces on a prey animal, striking it on impact to prevent
escape in thick vegetation. A single pounce may span l-4
meters and may be over a meter high (Geertsema 1985).
Another type of leap is vertical: birds and insects are seizedfrom the air by “clapping” the ‘front paws together(Smithers 1978) or striking with a downward
blow(Leyhausen 1979).
Geertsema’ s ( 1985) four-year study in the Ngorongoro
Crater is the most detailed investigation to date of serval
ecology. She found them to be largely crepuscular, restingin mid-day and occasionally at night. Females with kittens
increase diurnal hunting activity. Servals on farmland inSouth Africa’s Natal province were predominantly nocturnal, possibly a response to human disturbance (Bowland
1990). Through continuous observations (when possi-

Biology
Reproductive season: (W) Aseasonal, but birth peaks
appear to be correlated with wet seasons, when prey densities are at their highest due to new vegetative growth(Kingdon 1977, Smithers 1978). Geertsema (1985) suggests that a peak occurs in the mid- to late dry season in the
Ngorongoro Crater, so that post-rains high prey density
coincides with the raising of older but still dependent kittens.
Estrus: (C) 4 days (n=l: Mellen 1989).
Gestation: (C) 73 days (n=15; range 70-79) (Stuart andWilson 1988).
Litter size: (W) 2.5 (n=7; range l-3) (Smithers 1978); (C)1.96 (n=20: Skinner and Smithers 1990); 2.45 t 0.21
(n=14: Mellen 1989); range l-5 (Stuart and Wilson 1988).
Age at independence: (W) 6-8 months. Newly independent juveniles, tolerated by their mothers, may circulate
within their natal range for periods up to and over a year
(Geertsema 1985).
Age at sexual maturity: (C) 18-24 months (P. Andrews
litt. 1993).

inLongevity: (C) up to 19 years (Green 1991).
Habitat and Distribution
In sub-Saharan Africa, servals are found in well-watered
Savannah long-grass environments (Shortridge 1934,
Rosevear 1974, Smithers 1978), and are particularly associated with reedbeds and other riparian vegetation types(Geertsema 1985, Bowland 1990). This association withwater sources means that their distribution is strongly
localized over a wide area and within a variety of habitat
types (Fig. 5). They range up into alpine grasslands(Ansell and Dowsett 1988), up to 3,200 m in Ethiopia(Yalden et al. 1980) and 3,800 m in Kenya (York 1973).
Servals can penetrate dense forest along waterways andthrough grassy patches, but are absent from the rain forestsof central Africa. A few records from arid parts of southwestern Africa, Ethiopia and Somalia indicate that servals
will occasionally make use of sub-optimal habitats

22

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,ServalFrench hunter in 1936 in Arzew (northwest coast), said tobe the last in the area. There have been scattered reports ofserval occurrence throughout northern Algeria during the
1980s but zoologists have not been able to confirm them
(De Smet 1989, K. de Smet in Zitt. 1993). Surviving animals are likely to have been isolated from sub-Saharanpopulations for at least 6,000-7,000 years (Swift 1975).

(Shortridge 1934, Yalden et al. 1980, Stuart and Wilson
1988, A. Simonetta in Zitt. 1992).
In north Africa, relict populations may still be found in
humid scrub and mixed woodlands of Morocco’s Atlas
Mountains (Lambert 1966) and northern Tunisia andAlgeria (Gouttenoire 1954, De Smet 1989). The last confirmed record from Algeria is of an animal killed by a

Figure 5. Distributionof the serval (L. sewal).
1. Abuko IV (Gambia);
2. Mont Sangbe II; 3. Comoe II# (Ivory Coast); 4. “W” II* complex (Burkina Faso,
Benin and Niger); 5. Kwiambana
VIII complex; 6. Yankari II (Nigeria);
7. Zakouma
II complex (Chad);
8. Bamingui-Bangoran
II* complex (Central African Republic);
9. Simien Mts.** II; 10. Yangudi Rassa
II; Il. Bale Mts II; 12. Abijatta-Shalla
Lakes II (Ethiopia);
13. Boma II (Sudan);
14. ConkouatiIV;
15. Odzala II* complex (Congo);
16. Upemba
II; 17. Virunga II** complex;
18. Garamba
II** complex(Zaire); 19. Queen Elizabeth II* complex (Uganda);
20. Aberdare
II (Kenya);
21. Maasai Mara II (Kenya)+ Serengeti
II# (Tanzania)
complex; 22. Selous IV** complex (Tanzania);
23. Kasungu
II (Malawi);
24. Mana Pools II** complex (Zimbabwe);
25. Kaudom VIII (Namibia);
26. Moremi IV; 27. Chobe II(Botswana);
28. Kruger II complex; 29. St. Lucia IV complex;
30. Natal Drakensberg
IV; 31. unconfirmed observationof a serval by a farmer near George,
Cape province,
South Africa (M. Bowland in
litf. 1993); 32. Djurdjura
II; 33. El Kala V* (Algeria: De Smet 1989).

23

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,LeopardPopulation Status
Global: Category 4. Regional (sub-Saharan Africa):
Category 3. Regional (north Africa): Category 2(A).
IUCN: not listed. Smithers (1978) reviewed the serval’s
distribution and concluded that its range has remainedlargely intact, shrinking only in the extreme north andsouth due to habitat loss in the wake of increasing urbanization and changes in land use (C. Stuart in litt. 1993).
Possibly servals were never very numerous in northAfrica, and water sources in the region are likely to be
focal points of human use and settlement. However, servals are highly tolerant of agricultural development, which
fosters increased rodent densities, as long as there is sufficient water and shelter available (Bowland
1990).
Kingdon (1977) notes that the serval has adapted well tothe cultivation-fallow
mosaic that is widespread over the
moister regions of Africa. Degradation of forests to savannah in west Africa probably favors the species.
Geertsema (1985) found minimum home ranges in
Ngorongoro to be 11.6 km2 for one adult male and 9.5 km2
for one adult female over four years. The male’s homerange overlapped those of at least two adult females, while
the ranges of three adult females showed minimal overlap. Bowland (1990) found larger home ranges for servals on South African ranchland: 16-20 km2 for two adultfemales and 3 1.5 km2 for one male, monitored for 4-5
months during the spring and summer.

Myers 1986, Cunningham and Zondi 199 1; L. Gadsby, F.
Hurst in Zitt. 199 1, E. Abe in Zitt. 1993); they are frequently
marketed as “cheetah” or “leopard.” While the scale of theharvest and its effect upon populations is difficult to judge,
the pelt trade appears to be primarily domestic (especiallyfor ceremonial or medicinal purposes) or tourist-oriented,
rather than international commercial exports (WCMC
unpubl. data; see Table 1 in Part II Chapter 4). The serval’s localized distribution around water sources may
increase its vulnerability to hunting; it will also climb atree when chased by hounds (Stuart 1985).
Servals occasionally kill domestic poultry and only
rarely young livestock (sheep and goats): studies of theirdiet in farming areas in Zimbabwe (Smithers 1978) andSouth Africa (Lawson 1987) found no evidence that predation was a problem. Bowland (1990) pointed out thatproblem animals which raid chicken coops can be easily
live-trapped for translocation. Although 17% of Namibian
farmers who indicated that servals were present on theirland reported livestock predation, none took any control
measures (legally permissible), indicating that the problem
is not serious. For comparison, 36% of the farmers reporting stock predation by African wildcats took control measures (Joubert et al. 1982). The serval’s preference forrodent prey should actually benefit farmers: Geertsema(1985) calculated that an adult serval will eat some 4,000
rodents a year.

Protection
StatusCITES Appendix II. National legislation: not protected
over most of its range. Hunting prohibited: Algeria,
Botswana, Congo, Kenya, Liberia, Mozambique, Nigeria,
Rwanda, South Africa (Cape province only). Hunting regulated: Angola, Burkina Faso, Central African Republic,
Ghana, Malawi,
Senegal, Sierra Leone, Somalia,
Tanzania, Togo, Zaire, Zambia. No legal protection:
Benin, Cameroon, Ethiopia, Gabon, Gambia, Guinea
Bissau, Ivory Coast, Lesotho, Malawi, Mauritania,
Morocco,Namibia, Niger, South Africa,
Sudan,
Swaziland, Tunisia, Uganda, Zimbabwe. No information:
Burundi, Chad, Djibouti, Guinea (IUCN Environmental
Law Centre 1986, Smithers 1986, Hecketsweiler 1988).

Action Planning
Project 38.

Leopard, Panthera par&s(Linnaeus, 1758)
Other Names
Panther (English); leopard, panthere (French); Leopard,
Panther (German); leopardo, pantera (Spanish); nebr(Amharic: Ethiopia); eduka, ekun, ogidan (Ibo, Yoruba:
Nigeria); !‘hbm (Ju/hoan Bushman: Botswana, Namibia);ngoye, nze, goye (Kota, Fang, Kwele: Gabon); damissa(Hausa: west Africa); chui (Kiswahili);kwach (Luo:Kenya, Uganda); oluwaru keri (Maasai: Kenya, Tanzania);loli, mabiti, kweyi, mabilanga, moli, ka’u (Mbuti Pygmy
dialects: Zaire); nkewe, sinkwe z inqwe (Setswana:
Botswana); shabeel (Somalia).

Principal Threats
Wetland conservation is the key to serval conservation.
Wetlands harbor comparatively high rodent densities compared to other habitat types, and form the core areas ofserval home ranges (Geertsema 1985, Bowland 1990). Of
secondary importance is degradation of grasslands throughannual burning followed by over-grazing by domestic
hoofstock, leading to reduced abundance of small mammals (F. Hurst in Zitt. 1991, Rowe-Rowe 1992).
Trade in serval pelts has been reported from many
countries (Yalden et al. 1980, Sayer and Green 1984,

Description
and Behavior (Plate 1)
The leopard has the widest distribution of the wild cats, andshows great variation in appearance and behavior. In general, the coat color varies from pale yellow to deep gold or
tawny, and is patterned with black rosettes. The head,24

Part I: Species

lower limbs and belly are spotted with solid black. Coatcolor and patterning are broadly associated with habitat
type. Pocock (1932a) found the following trends in coloration for leopards in Africa: (1) Savannah leopardsrufous to ochraceous in color; (2) desert leopards-palecream to yellow-brownin color, with those from cooler
regions being more grey; (3) rain forest leopards-dark,
deep gold in color; (4) high mountain leopards-evendarker in color than 3. Black leopards (the so-called “black
panthers”) occur most frequently in humid forest habitats(Kingdon 1977), but are merely a color variation, not a subspecies. Variation in pelage has been the chief basis forthe description of numerous subspecies of leopard, 24 in
sub-Saharan Africa alone (Smithers 1975). However,Miththapala (1992), using molecular analysis and cranialmeasurements, concluded that sub-Saharan African leopards showed too little difference to warrant subspecificdivision and proposed that the 10 sub-Saharan subspecies
she examined should be subsumed into P.p. pardus, thenameoriginally appliedto the north African leopard.
The leopard is well known for its versatility asa generalist predator, and showsa number of morphological
adaptationsto this end, including its size, which showswide variation acrossits range. Exceptionally large males
weighing over 91 kg have been reported from South
Africa’s Kruger National Park (Turnbull-Kemp 1967),
where averageadult weightsare otherwise58 kg for males(n=3) and 37.5 kg for females(n=5: Bailey 1993). Maleleopards from the coastal mountains of South Africa’s
CapeProvince are much smaller,with an average weightof 3 1kg (n=27: Stuart 1981). Norton (1984) suggeststhat
this is becauseprey speciesare smallerin thesemountains.
In the rain forestsof northeasternGabon, one adult female
weighed 26 kg, and two malesweighed 34 and 41 kg (S.
Lahm in litt. 1993). In the rain forest of the Ivory Coast’s
Tai National Park, on the other hand, a male leopard was
captured which weighed 56 kg (Jenny 1993), and two
femalesweighed32 kg and 33 kg (Jenny in Zitt. 1994).
Despiteits relatively smallbody size,the leopardis still
capableof taking large prey. Its skull is massive,giving
ample room for attachmentof powerful jaw muscles. Its
whiskersare particularly long and there are often several
extra long hairs in the eyebrows, protecting the eyes andassisting movement through vegetation in darkness(Skinner and Smithers1990). Its scapulais adaptedfor theattachment of powerful musclesthat raise the thorax,
enhancing its ability to climb trees (Hopwood 1947).
Leopards can live independent of water for periods oftime, obtaining moisturerequirementsfrom prey (Bothma
and Le Riche 1986).
The known prey of the leopard rangesfrom dung beetles(Fey 1964)to adult maleeland(Kingdon 1977),whichcanreach900 kg (Stuart and Stuart 1992a). Bailey (1993)
found that at least92 prey specieshave beendocumented

Accounts.

Chapter

1. Sub-SaharanAfrica,Leopardin the leopard’sdiet in sub-SaharanAfrica. The flexibility of the diet is illustrated by Hamilton’s (1976) analysisof leopardscatsfrom Kenya’s Tsavo West National Park,of which 35% contained rodents, 27% birds, 27% small
antelopes,12% large antelopes, 10% hyraxes and hares,
and 18% arthropods. Seidensticker (1991a) and Bailey(1993)reviewed the literature, andconcludedthat leopards
generally focus their hunting activity on locally abundant
medium-sized ungulate speciesin the 20-80 kg range,
while opportunistically taking other prey. For example,
analysis of leopard scats from a Kruger NP study area
found that 67% containedungulateremains,of which 60%
were impala, the most abundant antelope, with adultweights of 40-60 kg. Small mammalremainswere foundmost often in scats of sub-adult leopards, especiallyfemales(Bailey 1993). Studieshave found averageintervals between ungulate kills to range from seven (Bailey1993)to 12-13days (Hamilton 1976,Le Roux and Skinner1989). Bailey (1993) estimatedaveragedaily consumption ratesat 3.5 kg for adult malesand 2.8 kg for females.
However, the leopard has an exceptional ability toadaptto changesin prey availability, and hasa very broad
diet. Small prey are taken where large ungulatesare lesscommon. For example, Grobler and Wilson (1972) and
Norton et al. (1986) analyzed leopard scatstaken from
Zimbabwe’sMatopos National Park and the mountainsof
southwesternCapeprovince and found rock hyraxes, common in the study areas,to be the most frequently taken
prey. In central African rain forest, both Jenny (1993) and
J. Hart and M. Katembo (in prep.) found the diet to consistmainly of duikers and small primates. Jenny (1993)
notes also that some individual leopards have shown astrong preference for pangolins and porcupines. In his
study area, the Ivory Coast’s Tai’ National Park, a longterm study of chimpanzeesdeterminedleopardpredation
to be the major causeof chimp mortality (Boesch 1991),
but D. Jenny (in Zitt. 1994) believes this may have been
the work of a specialistchimp-killing leopard. In the interior areasof South Africa’s Kalahari Gemsbok National
Park, where springbok are lessabundant,Bothma and Le
Riche (1984) found that 80% of leopard kills located bytracking (n=30) weighed lessthan 20 kg; nevertheless,
37% of all kills consistedof ungulates.By usingthe tracking method, they found that male leopards killed every
three days on average, and females with cubs every 1.5
days. At 3,900 m in the Kilimanjaro Mountains of
Tanzania, Child ( 1965)reported the leopard’sdiet to consist mainly of rodents, while Fey (1964) describeshow aleopard strandedon an islandin the wake of Kariba Dam
subsistedprimarily on fish (Tilapia), even though impalaand commonduiker were presentin low numbers.
The leopard shows several behavioral adaptations
which permit it to compete successfully with other large
predators, the first being its dietary flexibility. Bertram

25

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,LeopardLitter size: (C) 1.65 (range 1-4; n=59) (Eaton 1977); (W)(according to time of first observation, when cubs may be
several weeks old and some may have died) 2.13 (range
2-3; n=16) (Martin and de Meulenaer 1988).

(1982) studied radio-collared lions and leopards in thesame area in the northern Serengeti and found that, while
their ranges overlapped, leopards preyed on a wider rangeof animals than did lions, and there was little overlap
between their diets. Secondly, leopards often cache large
kills in trees. Great strength is required: there have been
several observations of leopards hauling carcasses ofyoung giraffe, estimated to weigh up to 125 kg (2-3 times
the weight of the leopard) up to 5.7 m into trees (Hamilton
1976, Scheepers and Gilchrist 1991). This behavior ismore common in areas where competing carnivores are
numerous (Schaller 1972, Bothma and Le Riche 1984);
where they are not, leopards may still drag the carcasses oflarge prey some hundreds of meters from the kill site into
dense vegetation or a rock crevice (Smith 1977). Leopards
may also retreat up a tree in the face of direct aggression
from other large carnivores. In addition, leopards havebeen seen to either kill or prey on small competitors, e.g.
black-backed jackal (Estes 1967), African wild cat (Mills
1990) and the cubs of large competitors (lion, cheetah,hyenas, wild dogs: Bertram 1982). Leopards have also
been observed to ambush terrestrial prey by leaping downfrom tree branches, although this behavior is apparently
opportunistic and relatively uncommon (Kruuk and Turner1967); like other cats, they probably generally prefer to gettheir footing on the ground before launching the actualattack (Leyhausen 1979). While the diet of rain forest
leopards may include arboreal animals (40% of scats from
Tai NP contained arboreal species, including seven speciesof primate: Hoppe-Dominik1984), they are unlikely toforage much in trees: radio-collared leopards in Tai’ have
only been observed to attack monkeys when on the ground(D. Jenny in litt. 1994).
Leopards are generally most active between sunset andsunrise, and kill more prey at this time (Hamilton 1976,
Bailey 1993). In Kruger NP, Bailey (1993) found thatmale leopards and female leopards with cubs were relatively more active at night than solitary females. The highest rates of daytime activity were recorded for leopards
using thorn thickets during the wet season, when impala
also used them (Bailey 1993). In tropical rain forest, D.
Jenny in Zitt. (1994) reports that two radio-collared leopards (an adult male and female) have hunted only during
the day, although they often travel at night.

Cub survival: (W) first-year mortality estimated at 41%(Martin and de Meulenaer 1988) to at least 50% annually(Bailey 1993).
Sub-adult survival: (W) Average annual mortality of subadults (1.5-3.5 years old) was estimated in Kruger NP at
32%, nearly twice as high as adults, probably related topoorer hunting success. Females: 40%; males: 25%(Bailey 1993).
Interbirth interval: (W) average 15 months (Martin andde Meulenaer 1988; these data include some shorter periods after litters did not survive) to over 2 years (Schaller
1972, Bailey 1993).
Age at independence: (W) 13-18 months (Bailey 1993,
Skinner and Smithers 1990). Siblings may remain together for several months before separating (Skinner andSmithers 1990). Dispersal may be delayed in areas whereprey are abundant, especially if adjacent habitat is occupied by resident leopards (Bailey 1993).
Age atfirst reproduction: females: (C) 33 months (range
30-36: Weiss 1952), (W) average 35 months (n=8: Martinand de Meulenaer 1988); males: 2-3 years (C: Green
1991).
Reproductive rate: (W) Bailey (1993) reported that the
average proportion of adult females producing young eachyear in his Kruger NP study area was 28%, while notingthat in some years no females gave birth, while in others up
to half of the females produced young.
Sex ratio of resident adults: (W) 1 male: 1.8 females(Bailey 1993, Hamilton 198 1).
Age at last reproduction: (C) average 8.5 years at one zoo(females: Eaton 1977), but up to 19 years (both sexes: A.
Shoemaker in Zitt. 1993).
Adult mortality: (W) average 19% annual mortality for
adult leopards in Kruger National Park. Old males 30%;
prime males 17%; old females 17%; prime females 10%.
The proportion attributable to starvation was 64% (Bailey1993).

Biology
Reproductive season: (W) probably year-round, but Bailey( 1993) found a peak in leopard births during the birth season of impala, the main prey species.

Longevity: (W) probably lo- 15 years (Turnbull-Kemp
1967, Martin and de Meulenaer 1988); (C) generally 12- 15
years, but up to 20 (A. Shoemaker in Zitt. 1993).

Estrus: (C) average 7 days.
Habitat and Distribution
Leopards occur in most of sub-Saharan Africa. They arefound in all habitats with annual rainfall above 50 mm

Estrus cycle: (C) average 46 days (Sadleir 1966).
Gestation: (C) 96 (90-105) days (Hemmer 1976).

26

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Leopardoccupies both rain forest and arid desert habitats. Leopards
range exceptionally up to 5,700 m, where a carcass was
discovered on the rim of Mt. Kilimanjaro’s Kibo Crater in1926 (Guggisberg 1975). They are abundant on the highest
slopes of the Ruwenzori and Virunga volcanoes, and havebeen observed to drink thermal water (37” C) in Zaire’s(Monod 1965), and can penetrate areas with less than thisamount of rainfall along river courses: e.g.leopards arefound along the Orange River in the Richtersveld National
Park (South Africa), which lies at the southernmost extension of the Namib Desert (Stuart and Stuart 1989). Out of
all the African cats, the leopard is the only species which

Figure 6. Distribution
and relative
abundanceof the leopard(I? pardus)
in sub-SaharanAfrica (after Martin and de Meulenaer
1988).
I. Niokolo-Koba
II# (Senegal);
2. Boucle du Baoule II complex (Mali); 3. Outamba-Kilimi
IV (Sierra
Leone); 4. Mt. Nimba I# complex(Guinea and Ivory Coast); 5. Sapo II (Liberia) + Ta’i II# (Ivory Coast)complex;6. Comoe II# (Ivory Coast); 7. Mole II (Ghana);
8. “W” II* complex (Burkina
Faso, Benin and
Niger); 9. Kainji Lake II (Nigeria);
IO. Zakouma
II complex(Chad); 11. Manovo-Gounda-St.
Floris II**(Central African Republic);
12. Southern
II; 13. Dinder II* complex (Sudan);
14. Simien Mts. II**;
15. Yangudi Rassa II (Ethiopia);
16. Dja IV# (Cameroon);
17. Lope IV (Gabon);
18. Odzala II* complex(Congo);19. Salonga II**, . 20. Upemba
II; 21. Virunga II** complex (Zaire); 22. Mt. Ruwenzori
II* complex (Uganda);
23. Tsavo II complex (Kenya);
24. Maasai Mara II (Kenya) + Serengeti
II# complex(Tanzania);
25. Selous IV** complex (Tanzania);
26. Nyika II (Malawi);
27. Zambezi
Wildlife Utilization
Area (Mozambique);
28. Hwange
II complex(Zimbabwe);
29. Kafue II complex; 29a. S. Luangwa
IIComplex(Zambia);
30. Kameia VI (Angola);
31. Etosha II (Namibia);
32. Central Kgalagadi
IIcomplex (Botswana);
33. Gemsbok
II (Botswana)+ Kalahari Gemsbok
II (South Africa) complex;
34. Richtersveld
V; 35. Cedarsburg
IV; 36. Kruger II complex(South Africa).

27

Pat? I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Leopardversality of the correlation of leopard density and rainfall;
and the desirability or not of re-opening commercial tradein leopard skins. R. Martin (in Zitt. 1994) concedes that avariable representing prey density should be incorporated
into the regression linking leopard density to rainfall.
Bailey (1993) also argues that while the link between herbivore density and rainfall may be generally valid, a herbivore biomass increase does not necessarily equate toincreased leopard prey biomass. The herbivore biomass
could be in the form of very large species (elephant, buffalo, hippopotamus) or herd-forming species (zebra andwildebeest), which provide little food for leopards.
Despite the controversy, there appears to be generalagreement that the leopard is not currently endangered in
sub-Saharan Africa, but that it is subject to local depletion
through exploitation and loss of habitat. Overall, Martinand de Meulenaer (1988) estimated the sub-Saharan population to number 7 14,000, based on their density/rainfall
regression. Although this figure is generally considered
to be an overestimate (Jackson 1989, Norton 1990), it represents the most practical and quantitative attempt to dateto estimate potential cat numbers across a large geographic
area. Its accuracy should be tested and improved by continuing investigation into leopard densities in key habitats, including tropical rain forest.
Biologists working in central African rain forest all
describe the leopard as common (M. Agnanga, R. Barnes,
A. Blom, J. Hart, S. Lahm in Zitt. 1993). The rainfall/density regression used by Martin and de Meulenaer (1988)
suggest that Zaire would hold some 33% of sub-SaharanAfrican leopards, a figure resulting from presumed very
high densities in tropical rain forest (up to 40 leopards,
including young and transients, per 100 km2). However,
Bailey (1993) is among several authorities who haveargued that since terrestrial mammalian prey biomass islower in rain forest than in Savannah environments, as thebulk of productivity is locked up in the tree canopy, therefore leopard density should be correspondingly lower.
Two studies are currently underway which should
eventually yield the first good data on leopard abundancein this habitat type (J. Hart, D. Jenny in prep.). D. Jenny(in Zitt. 1994) provides a preliminary estimate of five adultleopards in his 80 km2 study area in Tai’ NP, or 6.25 leopards per 100 km 2. J. Hart (in Zitt. 1994) offers a preliminary estimate of one adult leopard per 8- 12 km2 in Zaire’s
Ituri forest, or 8.3-12.5 leopards per 100 km’. These estimates are considerably lower than the 40 leopards per 100
km2 suggested by Martin and de Meulenaer’s rainfall/density regression. Yet they are also higher than adult leopard densities estimated for the Seronera woodland area of
Tanzania’s Serengeti NP (3.5 [Schaller 19721 - 4.7[Cavallo 19931 per 100 km2), which are among the greater
densities on the rainfall/density regression if the rain forest
estimates are excluded. In South Africa’s Kruger NP,

Virunga National Park (J. Verschuren in Zitt. 1993).
The leopard appears to be very succesful at adapting toaltered natural habitat and settled environments in theabsence of intense persecution. There are many recordsof their presence near major cities (e.g.Turnbull-Kemp
1967, Guggisberg 1975, Tello 1986a, Martin and deMeulenaer 1988: 18; G. Davies, B. Hoppe-Dominik,
R.
Kock, P. Norton in Z&t. 1993). Hamilton (1986b) reports
their occurrence in western Kenya in extensively cultivated districts with more than 150 persons/km2, the largest
livestock populations in the country, little natural habitatand prey, and where 20 years ago they had been considered extirpated.
However, leopards appear to have become rare throughout much of west Africa (Martin and de Meulenaer 1988:
1 I- 14). According to T. Anada (in Zitt. 1993), they havecompletely disappeared from much of the western Sahel.
Figure 6 shows the distributionof the leopard.
Countries are coded for abundance as determined byMartin and de Meulenaer (1988) (see explanation below),
except that equatorial Guinea, Mali, Nigeria, and Zimbabwe have been down-graded one category.
Population Status
Global: Category 5a(A). Regional: Category 4(A). IUCN:
not listed. The status of the leopard in sub-Saharan Africa
has been a matter of controversy since 1973, when it was
first listed on CITES Appendix I due to fear about the
impact of the then considerable international trade in leopard skins (Myers 1973). Six attempts have since been
made to determine the leopard’s status (Myers 1976, Teerand Swank 1977, Eaton 1978, Hamilton 198 1, Martin andde Meulenaer 1988, Shoemaker 1991). The first four
relied mainly on interviews
and questionaires, but
Hamilton’s (1981) work was more intensive, supplemented by the author’s own field studies, and focused
wholly on Kenya as a microcosm of the forces impacting
leopard populations throughout the continent. Martin andde Meulenaer (1988) also carried out wide-ranging interviews, but carried the process one step further by developing a population model for the leopard, which they usedin combination with a regression linking leopard densities
with annual rainfall to predict numbers of leopard in theregion. More recently, Shoemaker (1991) conducted an
extensive literature review and global correspondence tosummarize the status of the leopard throughout its entire
world range.
The first five studies were criticized from different
viewpoints (e.g.Hamilton 198 1: 93-94, USFWS 1982,
Martin and de Meulenaer 1988: xv-xx, Jackson 1989,
Norton 1990), with the debate focusing chiefly on theaccuracy of various population estimates; the model’s failure to account adequately for persecution and reductionof wild prey as factors lowering leopard density; the uni-28

Part I: Species

Bailey (1993) estimated average leopard density at 3.5
adults per 100 km”, with much higher densities of up to
30.3 per 100 km2 in the riparian forest zones, with highprey density. Leopard densities are lowest in arid environments (Martin and de Meulenaer 1988): for example,
1.25 adults per 100 km2 in South Africa’s Kalahari
Gemsbok NP (Martin and de Meulenaer 1988, based on
Bothma and Le Riche 1984). Hamilton (1981) andCavallo (1993) found that multiplying the number of adultresidents by 1.7 accurately accounted for the total numberof known animals in their study areas.
Leopards appear to be least numerous in west Africa,
possibly due to high levels of hunting for their skins, anddepletion of prey due to the trade in bushmeat (Myers
1976, Martin and de Meulenaer 1988). T. Anada (in litt.
1993) considers the leopard to be more rare than the lion in
the Savannah regions, while severely reduced abundance
was also reported from the west African rain forest zone
(Martin and de Meulenaer 1988). Also, in South Africa,
the leopard has been extirpated from many areas (Stuart
et al. 1985, Norton 1986, Rowe-Rowe 1992).
Leopard home range sizes determined by radiotelemetry have averaged between 30-78 km2 (males) and 15-16
km2 (females) in protected areas (Tsavo NP: Hamilton
1981; Kruger NP: Bailey 1993; Serengeti NP: Bertram
1982; Cedarberg Wilderness Area [South Africa]: Nortonand Henley 1987). Long-term observations of individual
female leopards have yielded larger estimates of homerange size in protected areas: 23-33 km2 (Le Roux and
Skinner 1989) and 37-38 km2 (Cavallo 1993). Bailey(1993) found the ranges of adult females were centeredon the most prey-rich habitat (riparian vegetation), while
the larger male ranges included lower quality habitat. In
mountainous terrain interspersed with farms and ranches,
Norton and Lawson (1985) found leopard home ranges of
338-487 km2 (for a male and female, respectively), suggesting both severely reduced prey availability and low
leopard density. On a Kenyan cattle ranch which maintained wild ungulates, Mizutani ( 1993) found female leopard home ranges to average 18 km2 (n=4) and males 55
km2 (n=4). In the TaYrain forest in Ivory Coast, Jenny (inZitt. 1994) reported a male range of 80 km2 and a female
range of 25 km”.

Accounts.

Chapter

1. Sub-SaharanAfrica,Leopardeither by landowners or government authorities, generally
permitted. Hunting prohibited or restricted to “problem/
dangerous” animals: Angola, Benin, Burkina Faso, Cameroon, Congo, Djibouti, Equatorial Guinea, Gabon, Ghana,
Guinea Bissau, Ivory Coast, Liberia, Mali, Mauritania,
Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia,
Sudan, Togo, Uganda, Zaire. No legal protection:
Gambia. No information: Burundi, Chad, Guinea (IUCN
Environmental Law Centre 1986, Shoemaker 1993).
Principal Threats
Although the leopard appears tolerant of habitat modification and occurs in the vicinity of settled areas, density
is certainly reduced when compared to occurrence in natural habitat (perhaps as low as l/10 or even l/100, as estimated by Martin and de Meulenaer [ 1988]), and theleopard becomes more vulnerable to exploitation and population fragmentation.
The fur trade was a major threat to the leopard in someareas during the 1960s and 1970s before the market collapsed due to changing public opinion and the impositionof international trade controls under CITES (see Part II
Chapter 4). Hamilton (198 1) reported that poaching forthe fur trade substantially reduced the leopard populationin Kenya, and considers the species to be particularly vulnerable to baited trapping, as leopards patrol small homeranges along regularly used trails. Use of poisoned baits is
also an important threat (Myers 1976; E. Abe, T. Anada, P.
Chardonnet, A. Simonetta in lift. 1993). Martin and deMeulenaer (1988) simulated the effects of high harvests on
leopards in east Africa during this period (they estimated
30,000 leopards killed between 1968-69), and concurredwith Hamilton’s (198 1) finding that hunting had severely
depressed populations there. However, their model also
indicated that even very high offtakes, of the order of
6 1,000 animals a year, had produced only a slight decline
in the total sub-Saharan population (see Part II Chapter
5). They consider the leopard to be generally resilient toharvest up to a critical threshold, which varies with density.
Martin and de Meulenaer (1988) argue that re-opening the fur trade with appropriate controls under CITES
would significantly benefit conservation of the leopard by
allowing local people to derive economic value from thespecies, seldom possible under current tourism and sport
hunting practices of most range states. Rural people areat present the force responsible for the continuing declineof the leopard in the region, through degradation of habitat where their livestock graze and persecution of the leopard as a threat to these animals. Development of options toenable local people to obtain income from leopards could
encourage them to refrain from eradicating the leopardsin their vicinity. Cobb (198 1), without considering such
options, could not foresee a future for the leopard in Africa
outside of protected areas. In 1986, protected habitat madeProtection
StatusCITES Appendix I. A system has been in place since 1983
by which selected African countries accept an annual
quota for the export of legitimate sport hunting trophiesand skins. As of 1994, the quotas are as follows: Botswana(130), Central African Republic (40), Ethiopia (500),
Kenya (80), Malawi (50), Namibia (loo), Mozambique(60), South Africa (75), Tanzania (250), Zambia (300),
Zimbabwe (500). National legislation: largely protectedacross its range, although killing of “problem” animals,

29

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Caracalsuccessful predation on adult springbok (Avenant 1993)
and young kudu (Shortridge 1934) has been reported.
After making a kill, caracals have been reported, leopard-like, to cache the remains in a tree (Roberts 195 1,
Mills et al. 1984), although this behavior is apparently
not common. Caracals have rarely been recorded to takecarrion (Skinner 1979, C. Stuart in Zitt. 1991). However,in Namibia’s Etosha National Park, a young adult female
scavenged from a springbok killed by a cheetah, waiting
for two hours for the cheetah to finish eating and moveoff (B. Bjil and K. Nowell pers. obs.). Moolman (1984a)
successfully
captured caracals for his study in the
Mountain Zebra National Park, South Africa, by placing
box traps near half-eaten large prey (mountain reedbuck)originally caught by a caracal, to which the animal eventually returned to feed. Shortridge (1934) states that they
are fairly easy to trap, as long as the bait is fresh. A female
with cubs was observed to return to feed on her springbok carcass for 3-4 consecutive nights (Avenant 1993).
Caracals are also known for their exceptional ability tocatch birds, leaping high into the air to knock them downwith their front paws. Avenant (1993) found that bird
remains occurred in 18% of caracal scats in the West
Coast NP, while Moolman (1984b) found their occurrence
in only 2-4% of scats collected in and around theMountain Zebra NP. Invertebrates and reptiles are also
eaten. Mean daily food intake for captive adult caracals
has been estimated by Moolman (1986) at 500 g for males
and 3 16 g for females. Caracals are predominantly nocturnal, but are often observed in the daytime, particularly
in protected areas.

up only 13% of potential leopard range (MacKinnon and
MacKinnon 1986a, Martin and de Meulenaer 1988).
Action Planning
Projects 33 and 34.

Caracal, Caracal caracal(Schreber, 1776)
Other Names
Desert lynx (English);caracal (French);caracal,Wiistenluchs (German); caracal, lince africano (Spanish);rooikat, lynx (Afrikaans: South Africa); delg ambassa(Amharic:
Ethiopia); djime taikorlo (Baguirmien);soumoli (Bornouan); guette anasa (Chad); filiki (Djerma);
pyaberi (Gourmanche: Burkina Faso); messo (Hausa:
Sahel); !hab (Hei//kum Bushman: Namibia); simbamangu(Kiswahili); =ui (Ju/‘hoan Bushman: Botswana, Namibia);
mwai (Luo: Kenya, Uganda); indabutshe, intwane(Ndebele: Zimbabwe);
ayuku (Ovambo: Namibia);
safandu (Peul/Foulbe); thwane (Setswana: Botswana);
hwang, twana (Shona: Zimbabwe); gedudene, maharra(Somalia); daga (Toucouleur: northwest Africa); ngada(Xhosa: South Africa).
Description
and Behavior (Plate 3)
A distinctive feature of the caracal is the black back of its
large ears (its name comes from the Turkish “karakulak”
or “black ear”). The ears are topped with black tufts about
4.5 cm in length (hence the other popular name, lynx,
although the caracal is not closely related to the lynxes).
The conspicuous ears are believed to play a role inintraspecific communication (Kingdon 1977). Caracals
are generally uniformly tawny-brown to brick-red in coloration, although black individuals have been recorded(Rosevear 1974, Guggisberg 1975). Caracals are thelargest of the African small cats: males can weigh up to
18 kg (average 13 kg, Cape Province, South Africa; n=61)
and females up to 16 kg (average 10 kg in Cape Province;n=41) (Stuart 1981).
Caracals prey on a variety of mammals, with rodents,
hares, hyraxes and small antelopes forming the major partof their diet (Smithers 1971, Grobler 1981, Stuart 1982,
Moolman 1986, Palmer and Fairall 1988) in many areas.
In South Africa’s West Coast National Park, near CapeTown, Avenant (1993) found that rodents were the mostcommon prey remains found in caracal scats, occurringwith 89% frequency. Antelope remains were more common than rodents in 194 stomachs collected from individuals killed as problem animals in Cape Province (Stuart
198 1). Caracals are capable of taking relatively large prey:

Biology
Reproductive season: (W) probably year-round (Bernard
and Stuart 1987, Avenant 1993).
Estrus: (C) l-3 days.
Length of estrus cycle: (C) 14 days (n=15).
Gestation: (C) 78-81 days (Bernard and Stuart 1987, P.
Andrews in Zitt. 1993).
Litter size: (C) 2.2 (range 1-4; n=15); (W) wild pregnant
females were also found to carry an average of 2.2 fetuses(range 1-3; n=22) (Bernard and Stuart 1987). The size offour litters in the West Coast NP also averaged 2.25 (range
1-3: Avenant 1993).
Age atjirst reproduction: (C) 12.5-15 months (males) and
14- 16 months (females); gametogenesis can occur somewhat earlier (Bernard and Stuart 1987, P. Andrews in Zitt.
1993).
Interbirth interval: (W) probably(Bernard and Stuart 1987).

30

one litter annually

Part I: Species

Chapter

1. Sub-SaharanAfrica,Caracalmore scrubby, arid habitats (Kingdon 1977, Yalden et al.
1980, Stuart 1984) (Fig. 7). They are not found in the tropical rain forests (Rosevear 1974). In South Africa, where
they are relatively abundant, they have been recorded(unusually) from the evergreen and montane forests of thesouthern Cape province (Stuart and Wilson 1988). In
Ethiopia, caracals range up to 2,500 m (and exceptionally
up to 3,300 m) in the Bale and Simien Mountains (Yalden

Age at last reproduction: (C) one female gave birth at 18
years.
Longevity: (C) up to 19 years (P. Andrews

Accounts.

in litt. 1993).

Habitat and Distribution
Caracals inhabit the drier Savannah and woodland regionsof sub-Saharan Africa, with a strong preference for the

Figure 7. Distributionof the caracal (C. caracal) in sub-SaharanAfrica.
1. Niokola-Koba
II# (Senegal);
2. Boucle de la Pendjari II* complex (Benin) + “W” II* complex (Benin,
Burkina Faso and Niger); 3. Air and Ten&e VIII (Niger); 4. Yankari II (Nigeria);
5. Benoue II* (Cameroon); 6. Fada Archei Fauna1 Reserve (Chad); 7. Dinder II* (Sudan); 8. Nechisar II (Ethiopia);
9. Marsabit II (Kenya);
10. Karamoja
N and S VI complex (Uganda);
11. Ruaha II complex (Tanzania);
12. Kundelungu
II complex (Zaire); 13. Lengwe II (Malawi);
14. Kafue II complex (Zambia);
15. lona
VI (Angola);
16. Gonarezhou
II (Zimbabwe);
17. Banhine NP (Mozambique);
18. ltala IV; 19. Weza
IV; 20. Storms River IV; 21. Kalahari Gemsbok
II (South Africa) + Gemsbok
II (Botswana)complex;22. Karoo II (South Africa); 23. Namib-Naukluft
II (Namibia).

31

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Africanwildcattotal of 2,800 caracals in 198 1 (Joubert et al. 1982).
However, control efforts thus far appear to have had little
effect on caracal populations (N. Fairall in Zitt. 1993).
Caracals typically re-colonize farming areas following
local extirpation (Visser 1978).
Hunting for skins and “luxury bushmeat” is reported
to be a threat in west and central Africa, where the caracal
is more sparsely distributed (F. Hurst in Zitt. 1991).

et al. 1980).
Field studies have been carried out only in South Africa
and Israel (for the latter, see species account under North
Africa and Southwest Asia). In South Africa, adult malehome range sizes in various study sites in Cape Province
have ranged from 3 l-65 km”, and females from 4-3 1 km2(Stuart 1982, Norton and Lawson 1985, Moolman 1986,
Avenant 1993).
Population Status
Global: Category 5b. Regional: Category 4. IUCN: notlisted. The status of the caracal is satisfactory in subSaharan Africa. It appears to be most abundant in South
Africa and Namibia, where its range is expanding (Stuart
and Wilson 1988, Rowe-Rowe 1992), possibly linked to
local extirpation of black-backed jackals by farmers(Pringle and Pringle 1979, Stuart 1982, H. Berry in Zitt.
199 1). In the Savannah regions of west and central Africa,
it is less common and patchily distributed in pockets ofdrier habitat (Kingdon 1977).

African wildcat, Fe/issilvestris, lybica group(Forster, 1770)
Other Names
Chat gante, chat sauvage d’Afrique (French); Falbkatze(German); gato mantes, gato silvestre (Spanish).
Sub-SaharanAfrica: Vaalboskat (Afrikaans:South
Africa); ye-dw dimmet (Amharic: Ethiopia); kongo diakouma, yacoumawara (Bambara); larrouye (Bornouan);batou ana guesh, guette (Chad); !ores (Hei//kum Bushman:
Namibia); ochwi, ochawhi (Herero: Namibia); nyau
(Kikuyu: Kenya); kaka pori, kimburu, kaka mwitu@Swahili); gamsi lala (Kotoko); /nua (Ju/hoan Bushman:
Botswana, Namibia); mbaki (Luganda); ogwang burra(Luo); igola (Ndebele: Zimbabwe); moula (Sara); phah,tib, (Setswana: Botswana); nhiriri (Shona: Zimbabwe);
wunndu ale (Wolof); mpaka, mbodla (Zulu: South Africa).

Protection Status
CITES Appendix II. National legislation: not protected
over most of its range. Hunting prohibited: Angola,
Benin, Burkina Faso, Cameroon, Ethiopia, Kenya,
Mauritania, Mozambique, Nigeria, Zaire. Hunting andtrade regulated: Botswana, Central African Republic,
Senegal, Somalia, Tanzania, Zambia. No legal protection: Congo, Gabon, Gambia, Guinea Bissau, Ivory Coast,
Lesotho, Malawi, Mali, Namibia, Niger, Rwanda, South
Africa, Sudan, Swaziland, Togo, Uganda, Zimbabwe.
Legal status as problem animal: Namibia, South Africa.
No information:
Burundi,Chad, Guinea (IUCN
Environmental Law Centre 1986).

North Africa and Southwest Asia: Qit berri, qit elghamli, qit wahsi (Arabic: Middle East); sooner mousch ormesch (Arabic: Sahara region); emschisch boudrar, akriw,
mousch abrani (Berber); biss burree (Saudi Arabia);
kadees el khala (Sudan); bisad car, jifa, mukulel dur,dinaad dur, dinad dibadeed (Somalia); tarda-tarhda,
arheda, aghda (Tamahaq).

Principal Threats
Caracals are often killed for suspected predation on small
livestock, although this appears to be a pervasive problem
only in South Africa and Namibia. Analyses of stomach
contents and scats from parts of South Africa outside theprotected areas system have found domestic stock to makeup a significant portion of the caracal’s diet, with estimatesranging from 17-55% in different areas (Pringle andPringle 1979, Bester 1982, Stuart 1982, Moolman 1986).
Brand (1989) found that reported annual small stock losses
to caracal ranged up to 5.3 animals per 10 km2. Large
numbers of animals are destroyed by farmers each year:
Stuart (1982) reports that an average of 2,219 animals
were killed annually in control operations in the Karooregion alone between 193 1- 1952. Brand (1989) surveyed
problem animal hunting clubs in Cape Province, and foundthat numbers of caracals reported killed or captured annually ranged from 0.02- 1.6/10 km2. Farmers responding toa government questionnaire in Namibia reported killing a

Description
and Behavior (Plate 3)
The wildcat has a very large geographic range, and varies
locally in appearance. In general, from north to south there
is a gradation of coat thickness, intensity of ground color,and amount of “tabby” markings (Robinson 199 1).
Pocock (195 1) recognized 26 subspecies. These subspecies are not considered in this document, which follows
the taxonomy of Weigel (1961) and Hemmer (1978a) inrecognizing four groups of FeZis silvestris: the forest cats(silvestris group) of Europe, the Caucasus and Asia Minor;
the steppe cats (ornata group) of south and central Asia
(see Eurasia); the tawny cats (lybica group) of Africa andthe Middle East; and F. s. catus, the domestic cat. The status of the Zybica group throughout its range is presented
here under the common name “African wildcat.”32

Pat? I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,Africanwildcatmer from September-March (Skinner and Smithers 1990).
In Saudi Arabia, Harrison and Bates (1991) report thecapture of a pregnant female in Oman in Feb. In the northern Sahara, breeding takes place from January-March(Dragesco-Joffe 1993).

The Zybicn group is the most widespread, and these catsdiffer from the European forms by their lighter build, lessdistinct markings, and thin, tapering tails. The African
wildcat is very similar in size and appearance to thedomestic cat, and the two can be difficult to distinguish in
the field. In southern Africa, males weigh an average of 5
kg (n=42), and females approximately 4 kg (n=36)(Smithers 1971, Stuart 198 1). The background color ofits coat ranges from reddish to sandy yellow to tawnybrown to grey, and is typically marked with faint tabby
stripes and spots. A characteristic feature of this group is areddish or rusty-brown
tint to the backs of the ears
(Skinner and Smithers 1990, Harrison and Bates 1991,
Dragesco-Joffe 1993).
Wildcats are primarily nocturnal, especially in very hot
environments or in proximity to settled areas, but are also
active in early morning and late afternoon. Studies haveshown rodents to be the major prey species throughout
southern Africa (Zimbabwe: Smithers and Wilson 1979;
Botswana: Smithers 1971; Karoo region and Central
Namib Desert: Stuart 1977; South Africa: Stuart 1982,
Palmer and Fairall 1988; Natal prov., South Africa: RoweRowe 1978; western Cape coast, South Africa: Avenant1993). This prey preference is presumably similar throughout their range (Rosevear 1974, Kingdon 1977, de Smet
1989, Harrison and Bates 1991). A variety of birds, reptiles, and amphibians are also taken, as well as other mammals, including young antelope (Smithers and Wilson
1979). Insects and arachnids, including solifuges and scorpions, are frequently taken, perhaps in relation to seasonalrodent scarcity (Smithers 197 1, Stuart 1977, Harrison andBates 199 1). Wildcats seldom scavenge carrion (Gasperetti et al. 1986, Skinner and Smithers 1990).
The African wildcat is generally recognized as theancestor of the domestic cat (Pocock 1907a). Unlike feral
domestic cats, which sometimes live in large groups or“colonies,” African wildcats are solitary. Liberg and
Sandell (1988) point out that domestic cats tend to formcolonies in the presence of clumped, rich food resources(such as garbage dumps), remaining solitary where prey ismore evenly and thinly distributed. It is interesting that in
captivity, female African wildcats have assisted mothers inprovisioning of young with food (Smithers 1983), a behavior observed in feral domestic cat colonies. However, preliminary results from a radiotelemetry study in Saudi
Arabia indicate that wildcats persisted in solitary habits
while feral domestic cats formed groups around a garbage
dump. This suggests that the domestication process may
be the most important factor underlying the sociality offeral cats (Macdonald et al. 199 1), perhaps leading to abroadening of the diet to include scraps and carrion.

Gestation: (C) 56-63 days (Green 1991).
Litter size: (W) 3.4 (n=7, range 2-5) (Botswana: Smithers1971); (C) 1-5.
Age at sexual maturity: (C) 11 months.
Longevity: (C) up to 15 years (Green 199 1).
Habitat and Distribution
The African wildcat has a very broad habitat tolerance. It
appears to be absent only from tropical rain forest: reports
from this habitat type may refer to domestic cats, or possibly to hybrids (e.g. a recent report from northwestern
Congo [M. Agnanga in litt. 19931). It is thinly distributed
throughout the Nubian, Saharan, and Arabian deserts,
where it is generally restricted to mountains and dry watercourses (Gasperetti et al. 1986, Kingdon 1990, Skinnerand Smithers 1990, K. de Smet in Zitt. 1993). Wildcats
range up to ~3,000 m in the mountains of Kenya, Ethiopia,
and Algeria (Kingdon 1977, Yalden et al. 1980, DragescoJoffe 1993, K. de Smet in Zitt. 1993).
Density is expected to vary widely with prey availability. Mendelssohn (1989) estimated a density of one individual per km* in open oak forest on hilly, rocky ground in
Israel. Fuller et al. (1988) reported the home range of amale African wildcat near Nakuru, Kenya as 4.3 km?
Population Status
Global: Category 5c. Regional (sub-Saharan Africa):
East):
Category 5. Regional (north Africa/Middle
Category 5. IUCN: not listed. While F. silvestris is themost abundant of the felids, widespread hybridization withdomestic cats is leading to the increasing rarity of purewildcats (see below).
Protection Status:
CITES Appendix II. National legislation: not protected
over most of its range. Hunting prohibited: Algeria, Israel,
Mauritania, Morocco, Mozambique, Niger, Nigeria,
Tunisia.
Hunting regulated: Angola, Burkina Faso,
Ghana, Senegal, Somalia, Tanzania, Togo. No legal protection: Benin, Botswana, Cameroon, Central African
Republic, Congo, Egypt, Ethiopia, Gabon, Gambia,
Guinea Bissau, Ivory Coast, Kenya, Lebanon, Lesotho,
Malawi, Mali, Namibia, Oman, Rwanda, Saudi Arabia,
Sierra Leone, South Africa, Sudan, Swaziland, Uganda,
United Arab Emirates, Zaire, Zambia, Zimbabwe.
No
information: Burundi, Chad, Djibouti, Guinea, Iraq,
Jordan, Libya, Qatar, Syria, Western Sahara, Yemen

Biology
Birth season: (W) in southern Africa, chiefly in the sum-33

Part I: Species

Accounts.

Chapter

1. Sub-SaharanAfrica,AfricanwildcatFigure 8. Distributionof the African
wildcat(E silvesfris,/yMcz~ group).
I. Namib-Naukluft
II; 2. Etosha II (Namibia);
3. Central Kgalagadi
II (Botswana);
4. lona VI (Angola);
5. N and S Luangwa
II complex (Zambia);
6. Kundelungu
II complex (Zaire); 7. Selous IV** complex;
8. Serengeti
II# complex (Tanzania);
9. Tsavo II complex (Kenya);
IO. Garamba
II** (Zaire); 11. Mago
II + Omo II complex (Ethiopia);
12. Manovo-Gounda-St.
Floris II** complex (Central African Republic);
13. “W” II* complex (Burkina Faso, Benin and Niger); 14. Bane d’Arguin II** (Mauritania).

Principal Threats
The primary threat facing the African wildcat throughout
its range is hybridization with domestic cats (see also discussions under Eurasia). Hybridization has been taking
place over a long period of time, particularly in the north of
its range where domestic cats arose thousands of years
ago. Mendelssohn (1989) believes that male feral cats
have a competitive advantage over male wildcats in accessto estrous females, due to both their larger size and their(IUCN Environmental Law Centre 1986; R. Daly, R.
Khan, I. Nader, A. Serhal, K. de Smet in Zitt. 1993).
Occurrencein Protected Areas
It is increasingly likely that pure strains of African wildcat will be found only in protected areas remote fromhuman habitation. Those areas which may possibly protect populations of African wildcats isolated from feral
domestic cats are marked with an asterisk in Figure 8.

34

Part I: Species

occurrence,in many places, at higher densities.
Hybridization in captivity has shown that distinctive characteristics of the African wildcat, such as its long legs andreddish-backed ears, are lost (Smithers 1983), althoughhybrids have been found with red-backed ears (M.
Lindeque, pers. ~0121111.1993).Smithers (1986) believes it
inevitable that hybridization “will lead to the virtual
extinction of the African wildcat as we know it at present.”
Feral cats are found throughout the wildcat’s range.
Smithers (1986) reports that, in South Africa, it is now
impossible to find pure wildcats anywhere in the vicinity
of settlements where there are domestic cats. Smithers(197 1) comments on hybrids found in Botswana withwhite legs and white patches on their bodies, and G. Mills(in Zitt. 199 1) reported destroying such a specimen in the
Kalahari at least 75 km from the nearest human habita-

Accounts.

Chapter

1. Sub-SaharanAfrica,Africanwildcattion. J. Gasperetti (in Zitt. 1993) reports that a geologist
found a litter of domestic cat kittens in the Rub el Khali(Empty Quarter: uninhabited sand desert of the southeastem Arabian peninusla), hundreds of kilometers from either
water or the nearest Bedouin encampment. Several breeding programs have been started to conserve pure strainsof wildcat in captivity, but the strongest hope for survivalin the wild of pure wildcats lies in controlling feral cat
numbers in remote protected areas.
Mendelssohn (1989) also attributes the rarity of
African wildcats in Israel to their susceptibility to feline
panleukopenia, transmitted by feral cats, which are generally resistant.
Action Planning
Projects 10, 15,43, and 89.

35

Part I: Species

Accounts.

Chapter

2. NorthAfricaandSouthwest

Asia

Part I
Species Accounts

Chapter 2
North Africa and Southwest

Box 1
Vulnerability

Asia

Index to Species of the Region (in order of vulnerability)

Species

Habitat Association
St [Mar] (Tot) Score

Asiatic lion, P. lea persica
Cheetah, A. jubatus
Set-vat, L. sen/aF
Leopard, P. pardus
Sand cat, F. margarita*
Caracal, C. caracal
Jungle cat, F. chausb
African wildcat, F.s. lybica groupa

N: I[O] (1)

N: 2 [2] (4)N: 3 [0] (3)B: 5 [3]N: 2 [I]B: 5 [3]B: 5 [3]B: 3 [4]

(8)
(3)
(8)
(8)
(7)

-1
-1

Geog.
Score
Range
(106km*)

Body
Size
Score

Total
Score

Rankings: 0.03

L -1
L -1
M 0
L -1
s +I
M 0
s +I
s +I

-3
-3
-2

1
l(A)
2(A)
WV

s:
S:
M:
M:
W:
M:
W:

-1

0
-1
0
0
0

1.02

0.27
3.74
5.40
7.06
5.80
8.70

-1
-1
-1
0
0
+I
0
+I

Key:
* All of this species’ range lies within the regiona See species account in Chapter 1, Sub-SaharanAfricab See species account in Chapter 3, Tropical Asia

Habitat Association
St = number of strong + significant habitats
N = Narrow (-1); B = Broad (0)[Mar] = number of marginal habitats(Tot) = total number of habitats

Geographic Range (in millions of km*)
S = Small (-1); M = Medium (0); W = Wide (+I)
Body Size
L = Large (-1); M = Medium(0); S = Small (+I)

(A) = Actively threatened

Regional Criteria
Habitat Association: Narrow = l-4 habitat types; Broad = 7-8 habitat types
Geographic Range: Small = cl million km*; Intermediate = 3-6 million km*; Wide = 7-9 million km*
Body Size: Large = 35-135 kg; Medium = 7-20 kg; Small = ~6.5 kg

See the Introduction

to the Species

Accounts

for explanationof the vulnerability

36rankingsystem(pp. 2-6).

-1
0
+I
+I

+2

4
WA)
5a
5b

Part I: Species

Asiatic lion, Panthera leo
persica (Meyer, 1826)

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Asiatic

lion1994). The record total length of a male Asiatic lion
(including the tail) is 2.92 m (Sinha 1987).
Mean pride size, measured by the number of adultfemales, tends to be smaller than for African lions: mostGir prides contain just two adult females, with the largest
having five (Walker 1994: 1S), compared to averages of 46 for African protected areas. However, despite the small
population size, individual animals are not well known;
future monitoring combined with molecular analysis ofrelatedness could show that what are currently identified as
separate prides consist instead of smaller foraging groups
from larger prides. Coalitions of males defend homeranges containing one or more groups of females, but
unlike African lions, Gir males generally associate with
their pride females only when mating or on a large kill. A
lesser degree of sociality in the Gir lions may be a functionof the smaller prey available to them: the most commonly
taken species (45% of known kills), the chital, weighs onlyaround 50 kg (Johnsingh and Ravi Chellam 199 1). The
larger sambar deer is also frequently taken (15% of known
kills), and may be preferred (Ravi Chellam 1993).
However, domestic cattle have historically been amajor component of the Gir lions’ diet (Pocock 1939a).
Livestock hair was found in 75% of over 1,800 lion scats
examined by Joslin (1973), and in 48% of those examinedby Sinha (1987). The wild ungulate prey base hasstrongly increased since the early 1970s (see below), andthis is reflected in a shift in the lions’ diet: recent analysis of over 3,000 scats showed that nearly 70% contained
hair of wild ungulates. A significant proportion of known
lion kills (30-35%) still consists of livestock, but this isprobably overestimated due to the relative ease of locating
livestock kills as opposed to wild ungulate kills (Ravi
Chellam 1993, Walker 1994: 11). Availability of livestock may also affect the loose sociality of Gir lions:based on 56 observations of lions at livestock kills, itappears that males prey on livestock to a greater extent
than females (Ravi Chellam 1993).

Other Names
Lion d’ Asie (French); Asiatische Lowe (German); leande Asia (Spanish); sinh, sawaj (Gujarati); sinh, sher, untia
bagh [camel tiger] (Hindi); hawaj (Maldhari); simha(Maldhari, Kannada); babar sher (Persian).
Description
and Behavior (Plates 1 and 8)
This Species Account primarily concerns the Asiatic lion
P.Z. persica, but reference must be made to the Barbary(north African) lion P.Z. Zeo, the nominate subspecies andthe lion that appeared in Roman circuses. There appears to
be no record of contiguous populations of the two subspecies in historic times. The Barbary lion is extinct inthe wild-thelast record being one shot in Morocco in1920 (Grzimek 1975). Some lions in Temara Zoo in Rabat
were identified in 1974 by Leyhausen and Hemmer(Leyhausen 1975) as having physical characteristics ofthe Barbary lion: very clear light iris, rather than brown;mane spreading behind the shoulders and covering the
belly right to the groin, high occiput (back of the head),short legs, and deep chest (W. York quoted in introduction to Leyhausen 1975) but none appeared absolutely
flawless (Leyhausen 1975). Attempts to establish a scientific breeding program have so far failed, although somezoos have bred specimens (W. Frey in Zitt. 1993)
Today, the only living representatives of the lions once
found throughout much of southwest Asia occur in India’s
Gir Forest. These Asiatic lions are genetically distinct
from the lions of sub-Saharan Africa, although the difference is not large, being smaller than the genetic distance
between human racial groups. Based on genetic distance,
the Asiatic lion is estimated to have separated from theAfrican population as recently as 100,000 years ago, notlong enough for reproductive incompatibilities to have
evolved (O’Brien et al. 1987b,c). The most striking morphological character, which is always seen in Asiatic lions,but rarely in African lions, is a longitudinal fold of skinrunning along its belly (O’Brien et al. 1987~). In addition, male Asiatic lions have only moderate mane growthat the top of the head, so that their ears are always visible,
while many African males develop full manes which completely obscure the ears. Finally, about 50% of Asiatic lion
skulls from the Gir forest have bifurcated infraorbital
foramina (small apertures which permit passage of bloodvessels and nerves to the eyes). In African lions, there is
only one foramen on either side (Pocock 1939a, O’Brien etal. 1987~). Asiatic lions are slightly smaller than African
lions: adult Gir males weigh 160- 190 kg (n=4), whilefemales weigh 1 lo- 120 kg (n=2) (Ravi Chellam in Zitt.

Biology
Reproductive season: year-round, but based on sightingsof cubs, there is a birth peak from late winter to early summer (February-early April: Ravi Chellam in Zitt. 1994).
Litter size: (W) mean 2.5, range l-5 (observed only after
young cubs are fully mobile) (Walker 1994: 18); (C) 2-6(Chavan 1993).
Age at first reproduction: (W) field workers estimate
females 4 years, males 5-8 years; (C) 3 years (males andfemales) (Walker 1994: 18).
Age at last reproduction:(W) females 15-16 years(Chavan 1993); (C) both sexes 15 years (Walker 1994:
18). Adult sex ratio: 1 male:2.2 females (Ravi Chellam37

Pat? I: Species

Accounts.

Chapter

2. Norlh

Africaand Southwest

Asia,

Asiatic

lion

livestock live in the wildlife sanctuary which surrounds
the core national park. Within a 10 km radius surrounding
the sanctuary boundary, there is a human population of
160,000 and about 100,000 head of livestock (Walker
1994: 13-14). During drought years in the past, cattle
have been brought to graze in the protected area from hundreds of kilometers away, with numbers reaching up to
70,000 (Berwick 1976); the average annual number ofseasonally grazing livestock in the park is currently estimated at 20,000 (Walker 1994: 14).
The pastoralist Maldharis, who make up about onethird of the reserve’s human population, have been part of
the Gir ecosystem since approximately 1860 (Berwick1976). Their primary means of subsistence is selling ghee(clarified butter used for cooking). However, livestock
overgrazing has led to soil impaction and erosion, as wellas xerification of the forest. Berwick (1976) found that,
contrary to popular assumption, overgrazing was not leading to a decline in wild ungulate populations. Wild ungulates were found to feed mainly on woody plants rather
than grasses, and it was concluded that lion predation was
the primary factor limiting their numbers, then estimated at
6,200 (Berwick 1974). However, despite Berwick’s findings, with the removal of some 845 Maldhari familes andtheir herds, wild ungulates have greatly increased, and are
currently estimated at 43,000, including some 38,000 chital (Rashid 1984, Khan et al. 1990, Ravi Chellam andJohnsingh 1993a).

and Johnsingh 1993a). In captivity, records from the
Sakkarbaug Zoo, which maintains the largest captive population of Asiatic lions, also show a female bias (1 female:
0.38 male), but it is not known whether this is a local
effect, or whether it is representative of wild conditions(Walker 1994: 18).
Juvenile mortality: (< 12 mos) (W) 33% (3 cubs of 9 from
4 litters); (C) 36% (74 of 205 cubs born at Sakkarbaug
Adult mortality: (W) estimated at 8-lo%, based on anaverage of 10 adult animals per year which are removed
from the Gir population for health reasons and taken to
the Sakkarbaug Zoo (Walker 1994: 18).
Longevity: (C) females 17- 18 years but up to 2 1; males 1616 years but up to over 18 (Chavan 1993).
Habitat and Distribution
The range of the lion in north Africa and southwest Asia
formerly stretched across the coastal forests of northern
Africa and from northern Greece across southwest Asia
to eastern India (Guggisberg 1961, Joslin 1973, Smithers1975). It became extinct in eastern Europe around A.D.
100, and in Palestine around the time of the Crusades(Guggisberg 196 1). It remained widespread elsewhere
until the mid- 1800s when the advent of firearms led to its
extinction over large areas. By the late 1800s the lion
had disappeared from Turkey @stay 1990); the last reports
from Iran and Iraq date to 1942 (Joslin 1973) and 19 18(Hatt 1959) respectively. In India, lions ranged east to thestate of Bihar, but declined under heavy hunting pressure-Pocock(1939a) uses the diaries of an English officer who shot 300 lions during the 1857 Indian Mutiny as
an example. By the turn of the century, the Asiatic lion
was confined to the Gir Forest, where it was protected by
the Nawab of Junagadh in his private hunting grounds(Kinnear 1920).
The Gir is dry deciduous forest dominated by teak, thepredominance of which is partially due to the silvicultural practices of the Gujarat State Forest Department,
which permits logging and replants clear-cut areas withteak (Berwick 1976). The drier eastern part of the Gir isvegetated with acacia thorn Savannah and receives about
650 mm annual rainfall; rainfall in the west is higher at
about 1,000 mm a year (Ravi Chellam and Johnsingh
1993a). The forest, which covered about 2,600 km2 at theturn of the century (Oza 1983), has since shrunk to lessthan half this size. Most of the remaining forest is
included in the Gir National Park and Wildlife Sanctuary(259 + 1,153 = 1,412 km2). The Gir Forest is the last representative block of the natural vegetation of the semi-arid
Saurashtra peninsula, and is surrounded by cultivation.
Moreover, about 7,500 people and their 14,000 head ofPopulation Status
Global: Category 3(A). Regional: Category 1. IUCN:
Endangered. The Gir lion population had been reduced
to a very low number by the early years of the 20th century: fewer than 20 according to the Chief Forester ofJunagadh (Winter-Blyth1949). However, Gee (1964)
reported the “certainty” of the neighboring ruler, the Jam
Saheb of Nawanagar, that there were about 100 lions, andthat the ruler of Junagadh gave low numbers in order todissuade would-be trophy hunters.
The first census, calculated on the basis of individually
recognizable pug marks, was conducted in 1936, andyielded an estimate of 234 adults (Winter-BlythandDharmakumarsinhji
195 1). Subsequent censuses, basedon counts of animals at live buffalo baits, estimated the
population at around 100 adults between 1968-1979.
Censuses based on counts of animals both at waterholes
and baits conducted in 1985 and 1990 indicate that the
population is increasing, with 191 adults (66 males, 75
females, 50 sub-adults) counted in 1985, and 221 adults(99 males, 122 females, sub-adults not distinguished) in1990. In addition, about 30-40 lions are believed to livein the agricultural mosaic surrounding the reserve boundaries (Chavan 1993, Walker 1994: 5). The accuracy of
the waterhole count technique has been questioned (Kunte

38

Part I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Asiatic

lion

Boundariesof thelion’s historical range#
cl:igureProtected
remainingarea where only
population
occursPotential

re-introduction

1. Past and present

sites

distributionof the lion (R lea) in northAfricaand southwest

Asia.

Historical
range: Source is Guggisberg(1961) unless stated otherwise.
a. Aristotle and Herodotus
wrote that lions were found in the
Balkans in the middle of the first millenium
B.C. When Xerxes advanced
through Macedonia
in 480 B.C., several of his baggage camels
were killed by lions. Lions are believed to have died out within the borders of present-day
Greece in A.D. 80-100. b. Lions were probably
found in the Azerbaijanarea up to the 10th century A.D. Their disappearance
from the reed thickets and pistachio and juniper forests isprimarily associatedwith an increase in human population
and a change in environmental
conditions,which in turn led to the decline ofungulatesin the region (Heptnerand Sludskii 1972). c. Lions could still be found in the vicinity of Samaria in the 12th century.
d. Lions
disappeared
from the Moroccan
coast by the mid-l 800s. They may have survived
in the High Atlas Mountains
up to the 1940s. e. Last
known lion in Algeria killed in 1893 near Batna, 97 km south of Constantine.
f. Last known lion killed in Tunisia in 1891 near Babouch,between Tabarka
and Ain-Draham.
g. Lions were extirpatedfrom Tripolitania
as early as 1700. h. Last known lion in Turkey killed in 1870
near Birecik on the Euphrates@stay 1990). i. Sir Alfred Pease reported that lions still existed west of Aleppo, Syria, in 1891 (Kinnear
1920). j. Lions occurredin the vicinity of Mosul, Iraq, in the 1850s. The Turkish governor’s
bag of two in 1914 is the last report of themfrom the area (Kinnear 1920). k. Lions were reported to be numerous
in the reedy swamps bordering
the Tigris and Euphrates
rivers in
the early 1870s. The last known lion in Iraq was killed in 1918 on the lower Tigris (Hatt 1959). I. The valley of Dashtiarjan,
57 km west of
Shiraz in Iran, was famous for its lions in the late 1800s. m. The last known report of lion presencein Iran was a 1942 observationof apair near Dizful, by American
engineersbuilding a railway (Heaney1943). n. There are no confirmed
modern records of lion presenceincentral or eastern Iran, nor Afghanistan
or Baluchistan.
o. The last known lion in Pakistan killed near Kot Deji in Sind province in 1810. p.
However,a British admiral travellingby train reported seeing a maneless lion near Quetta in 1935, eating a goat: “It was a large lion, verystocky, light tawny in colour, and I may say that no one of us three had the slightest doubt of what we had seen until, on our arrival at
Quetta, many officers expressed
doubts as to its identity, or to the possibility of there being a lion in the district”.
q. The lion’s range may
have extended
as far east as Bihar and Orissa states: a lion was reportedly
killed in the district of Palamau (Bihar) in 1814. r. Last lion
recorded from the southernend of its Indian range killed at Rhyl in Damoh district, near the Narmada
river, in the cold season of 18471848 (Kinnear 1920). s. Fifty lions were killed in the district of Delhi between
1856-1858.
Twenty-fiveyears later Blanford (1891) wrotethat “in India the lion is verging on extinction.”
Present
range: 1. Gir II complex.
tion in the 1960s: 4. Chandraprabha

PotentialIV.

reintroduction

sites: 2. Palpur

39

Kuno (Kuno)

IV; 3. Sitamata

IV. Site of unsuccessful

reintroduc-Part I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Asiatic

lion

cumbersome and unrealistic (Joslin 1984, Ravi Chellam
and Johnsingh 1993a), and there are recent reports of villagers killing lions.
Even more alarming, the lions which have long been
famed for their docility toward humans have recently
begun to attack people, mainly during sorties outside thesanctuary. Saberwal (1990) has documented 8 1 attacks
resulting in 16 deaths from January 1988-April 1990, as
compared to 65 attacks resulting in eight deaths over theprevious decade. He suggested that the spate of attacks
was attributable to reduced availability of livestock prey
due to the effects of a severe drought in 1987-1988, andnoted that the attacks were clustered near 1) high humanpopulation density and 2) sites where lions were baiteduntil 1987 to show them to tourists. Lions in these areas,familiar with large groups of people, would have been lesssensitive to human threats, and thus more likely to havebecome involved in conflicts over livestock.
Ravi
Chellam and Johnsingh (1993a) stress that greater involvement of the impoverished Maldharis and villagers in andaround the Gir in the management of the protected area isa matter of highest priority.
The Asiatic lion currently exists as a single population, and is thus vulnerable to extinction from unpredictable events, such as an epidemic or large forest fire.
However, it is also a large, healthy population, and a recent
Population and Habitat Viability Analysis (PHVA) workshop in India (Walker 1994) predicted a zero percent
chance of extinction over the next 100 years, based on their
population model.
Nonetheless, establishment of at least one other wild
population is advisable for population safety, for maximizing genetic diversity, and in terms of ecology (re-establishing the lion as a component of the fauna in its former
range). The Asiatic lion PHVA (Walker 1994) reviewed
several potential translocation sites for suitability in termsof habitat and prey base, and selected the Palpur-Kuno
Wildlife Sanctuary in northern Madhya Pradesh as themost promising (this and other potential sites are shownas stars on the distribution map). The size of the protectedarea is currently only 345 km2, but it could be expanded
to approximately 2,000 km2 if adjacent forest were incorporated. Human disturbance is considered to be relatively
low-although
there are still 13,000 people and 16,000
livestock in the proposed area. Moving them out, as wasdone in several Tiger Reserves, would no doubt be
extremely difficult. Moreover, there is considerable hostility to wildlife in rural India, and moving lions into anarea where people have had no experience of them for generations is risky, both for the lions themselves and for thelarger cause of big cat conservation. A previous attempt toestablish a second population in the Chandraprabha
Wildlife Sanctuary in eastern Uttar Pradesh appeared to be
succeeding, as the population grew from three to 11 ani-and Gore 1986). The most reliable method would be tomark individuals, which could lead to improved understanding of population dynamics (Walker 1994: IO).
Radio-telemetry studies (Ravi Chellam 1993) estimate
the mean annual home range of male lions at 110 km2,
and females at 50 km? The ranges of male coalitions are
between 1OO-150 km2 in size, while those of single males
are smaller at 50 km2 (Chavan 1993). Density is estimatedat one lion per 7 km2, which would yield a population of
202 adults, a total very close to the 1990 census result(Ravi Chellam 1993). This density is comparable to theupper range of estimates of lion density in sub-SaharanAfrica. For comparison, tiger densities in good habitatwith abundant prey and low numbers of interspecific competitors (Kanha National Park, India and Royal Chitwan
National Park, Nepal) are of the order of one tiger per 1l17 km2 (Schaller 1967, Sunquist 198 1, Smith 1984,
Karanth 1987). The number of lions appears to haveexceeded the estimated carrying capacity of 200-250 animals (Ravi Chellam 1987, Rashid 1991, Chavan 1993,
Walker 1994: 5).
Genetic studies (O’Brien et al. 1987b) of 28 lions from
India’s Sakkarbaug Zoo (four wild-born founder animals
and 24 offspring of nine, including the four sampled, original founders) revealed total genetic uniformity among
the animals, similar to that found for cheetahs (O’Brien etal. 1986). A high incidence of spermatozoa1 abnormalities
has also been found for both wild and captive Asiatic lions(O’Brien et al. 1987c, Wildt et al. 1987b, Fouraker and
Wildt 1992). No signs of compromised reproduction in
the wild have been reported (P. Jackson, pers. comm.),but Walker ( 1990, 1994) has noted high rates of infantmortality among the inbred Sakkarbaug Zoo lions. On
the other hand, hybrid African-Asiatic lions breed well in
captivity (O’Brien et al. 1987~).
Protection Status
CITES Appendix I. National legislation: fully protected inIndia (Ravi Chellam and Johnsingh 1993b).
Principal Threats
The close proximity of predators, livestock, and humans in
the Gir Forest gives rise to a number of management problems which threaten the Asiatic lions. There are four large
temples located in the Gir Forest, which is cut by five
major roads and a railroad, so that a considerable volumeof people moves through the protected area. Lopping oftree branches for firewood is widespread, and is having adevastating effect, especially upon river-me forest, which isprime habitat for lionesses with cubs during the dry season(Ravi Chellam 1993). Lions have been preying on cattle
ever since they first moved into the area, but there are indications that peoples’ tolerance of lions is coming to an end.
The government’s livestock loss compensation scheme is40:Part I: Species

mals, but then the lions disappeared, presumably shot or
poisoned (Negi 1969).
Theoretically, the captive population of Asiatic lions canbe considered to represent a second population. A Species
Survival Plan (SSP) was established by the American Zoo
and Aquarium Association (AZA) to manage the >200
Asiatic lions held by western zoos. However, not only is
this SSP-managed population entirely descended from fivefounder animals, but two of the founders were African or
African-Asiatic hybrids, as demonstrated by genetic studies
and morphological characteristics (O’Brien et al. 1987~).
Only three individuals in North American zoos are of pure
bloodline (Wildt et al. 1992a). The total global captive
population of pure Asiatic lions is believed to be 82, of
which 23 are held outside of India (Walker 1994: 2 1). The
government of India is currently considering offering problem wild lions to western zoos as new founders. The
AZA’s Felid Taxon Action Group has recommended thathybrid lions may continue to be bred to monitor their vigor
until such time as space is required for pure Asiatic lions. It
also called for collection of germ plasm from wild animals,
which could be used to infuse genetic diversity into the captive population (Wildt et al. 1992a: SO).

Chapter

2. NorthAfricaand Southwest

Asia,

CheetahAsian cheetahs to differ in morphology (Hemmer 1988)
and pelage (pale fawn as opposed to sub-Saharan yellow,
with spots more widely spaced: Heptner and Sludskii
1972, C. Groves in Karami [1992]). Dragesco-Joffe(1993) has observed that cheetahs of the open, sandy
Saharan desert tend to be pale, with ochre rather than blackspots, and muted “tear line” and tail rings. There is a rare
form, locally called “white cheetah,” which is exceptionally pale. However, cheetahs living around the black rocksof the Saharan mountain ranges tend to retain the blackspots common to sub-Saharan cheetahs. Dragesco-Joffe
has also reported that Saharan cheetahs tend to be rather
small: two adult males killed in the Ten&e region of Niger
had a shoulder height of only 65 cm, as compared to 85 cm
for sub-Saharan cheetahs (Bowland et al. 1993). The
genetics of north African and southwest Asian cheetahs
have yet to be investigated.
While the question of evolutionary relationships
remains to be resolved, the main difference between cheetahs of this region and those south of the Sahara is that they
are much more rare. Some of this rarity is natural, given
the harsh conditions of sand desert. However, severe
depletion of the cheetah’s ungulate prey base (East 1992a,b) and direct persecution are the major threats to the cheetah’s survival in this region.
There is little information available on the ecology ofthese cheetahs. Gazelles are generally indicated as themain prey species (Heptner and Sludskii 1972, Harrisonand Bates 1991). In India, cheetahs took primarily blackbuck antelopes and chinkara gazelles, but were also
known to attack nilgai antelope and domestic goats andsheep (Pocock 1939a). In Turkmenistan, cheetahs primarily took goitered gazelles, and their disappearance
from this area is strongly associated with the decline ofgazelles in the mid- 1900s (Heptner and Sludskii 1972). In
Iran, cheetahs outside protected areas with gazelle populations are reported to prey mainly on hares, an abundantfood source because they are not usually taken by Muslim
hunters (M. Karami in Zitt. 1990). Cheetahs in subSaharan Africa are known to take hares opportunistically.
Whether cheetahs can subsist almost entirely on small
prey needs to be investigated.
Dragesco-Joffe (1993) reported that cheetahs living in
the Saharan mountains often hunt at night, when temperatures are cooler. He translates the Touareg name for cheetah as “one who advances slowly”-areversal of the
popular perception of the cheetah as one of the fastest landmammals. The name is a tribute to the cheetah’s slow,
patient stalking of gazelles in open terrain with very little
cover. Dragesco-Joffe also states that Saharan cheetahs
occasionally take ostrich and Barbary sheep.
Throughout this region and in Europe as well, captive
cheetahs were kept by the nobility and trained to hunt, apractice dating back about 5,000 years to the Sumerians.

Action Planning
Project 35.

Cheetah, Acinonyx
Schreber, 1776

Accounts.

jubatus

Other Names
Guepard (French); Gepard (German); guepardo, chita(Spanish); fahad (Arabic); yeoz (Brahui: Pakistan); pulam(Bukharian & Turkmenian); chita, laggar (Hindi: India);tazy palng (Dari: Afghanistan); yuz, yuz peleng (Farsi:Iran); ala bars, pyestrai, or pyatnistai bars (Kazakh); gurk
(Mekrani: Pakistan); tazy prang (Pashto: Afghanistan);Asiaskii gepard (Russian); addle amayas (Tamahaq,
Tamacheq [Touareg]): Northwest
Sahara); myallen,
koplon (Uzbek).
Description
and Behavior (Plate 8)
See full species account under Sub-Saharan Africa. Someauthorities consider the cheetahs of north Africa and southwest Asia to be a single race, A.j. venaticus (Pocock 1939a,
Ellerman and Morrison-Scott
195 1), while others argue
that north African populations have only become isolated
from populations at the southern edge of the Sahara within
the last century (K. de Smet in Zitt. 1993). Harrison andBates (199 1) label the distinction between Asian and
African cheetahs dubious, while other anatomists consider

41

Part I: Speciesq.

-igureAccounts.

Speciespresent

Historical

range

Chapter

2. NorthAfricaand Southwest

Asia,

Cheetah

range

0
A

Unconfirmed

recordProtected area where
species occurs

2. Past and present

distributionof the cheetah(A. julmtus)

in northAfricaand southwest

Asia.

Historical range: a. The Azerbaijankhans and Armenianand Kat-tlian (eastern Georgian)princes hunted with trained cheetahs
up to theChronicles(Kartlis
14th century.
In 1472, Josef Barbaro saw the “100” hunting cheetahs of an Armenian
prince. The GeorgianTskhovreba)
place the cheetah in eastern Georgia in the Middle Ages. Fossil remains dating to the middle Pleistocene
documentthecheetah’s presencein the Caucasus
region, but it is unclear whether wild cheetahs persisted there in historical times (Vereschagin1959).
b. Tristram (1866, cited in Harrison and Bates 1991) noted the presenceof a few cheetahs in Gilead, the vicinity of Mt. Tabor and the hillsof Galilee, but cheetahs have been extinct in this area for over 100 years (Harrisonand Bates 1991). c. Cheetahs
were still found up to 40
years ago in the Atlas mountainsof Morocco (Wrogemann1975). d. The last record for the cheetah in Western Sahara dates to when an
animal was captured
in 1976 and given to the Algiers Zoo. e. The last known cheetah in Tunisia was killed in 1960 near Bordj Bowrgiba
inthe extreme south of the country.
f. The last observationof a cheetah in Libya was in 1980 in the southwesternpart of the country bordering Algeria, where cheetahs are still known to exist (K. de Smet, pers. comm. 1990, cited in Kraus and Marker-Kraus
1991). g. Hardy(1947) mentions seeing two cheetahs in the Sinai Desert in 1946. h Last record of the cheetah in Yemen dates to an observation
by J.T.
Ducker in 1963 in Wadi Mitan (Harrisonand Bates 1991). i. Last known cheetah in Oman shot near Jibjat, Dhofar in 1977 (Harrison1983). j. Dickson (1949) remarked
on the presence of cheetahs
in Kuwait.
k. Cheetahs
were reported to be rare in the desert west of
Basra, Iraq, in 1926 (Corkill 1929). I. Last record of the cheetah in Iraq is a photographof one killed by a car between the HI and H2
pumping stations (Harrisonand Bates 1991). m Cheetahs
were killed in the early 1950s by oil workers
near the Saudi Arabian, Jordan
and Iraq border intersections(Hatt 1959). n. Last record for the cheetah in Saudi Arabia dates to 1973, when two were killed near Ha’iland exhibited for a few days near the lmara palace (Nader 1989). o. The last record of the cheetah in India, where the species was formerly widespread,dates to 1947, when the Maharajahof Korwai (misprinted
as “Korea” in J. Bombay /Vat. Hist. Sot. Vol. 47:719) innorthern Madhya
Pradesh, shot three cheetahs(with two bullets) at night, spotlighting
them with his car headlights.
Taxidermistsvanlngen and van lngen (1948) transmitted
the “record of this shoot” in a letter to the Journal of the Bombay Natural History Society.
The
editors appended
a note saying, “The editors were so nauseatedby the account of this slaughter that their first impulse was to consign itto the waste-paper
basket.
Its publication
here is intended in the nature of an impeachment
rather than any desire on their part to condone
or extol the deed.” p. Cheetahs
formerly occurred throughout
the dry hills west of the lndus river in Pakistan at the end of the 19th century,
but subsequent
reports are sparse and they are probably
now extinct (T. Roberts in litt. 1993). The last record is of a trade skin obtained
Continued

42

on next pagePat-t I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Cheetahin 1972, which reportedly
originatedfrom the Mekran border region near Iran (Roberts
1977, Groombridge1988). q-r. Habibi (1977) and
Sayer and van der Zon (1981) believe the cheetah to be extinct in Afghanistan,where it was formerly
found throughout
the lower steppes
up to 1,000 m. Skins were purchasedin fur markets in Fara (q) in 1948 and in Herat (r) in 1971, but their origin is not known.
s. Thecheetah has disappearedin recent times from the trans-Caspian
region (Bannikov
and Sokolov 1984). It was probably extirpatedfromthe Kyzylkum
desert region southeastof the Aral Sea in the early 196Os, and from the Ustyurt and Mangyshlak
regions west and southwest of the Aral by the late 1970s (Ishadov
1992; E. Matjuschkin,
E. Mukhina
in litt. 1993). The last unconfirmed
observationof a cheetah
in this region dates to 1982 on the Turkmenistan-Kazakhstan
border (s); the last confirmed
evidence of a small, establishedpopulation
dates from 1973 in Turkmenistan,further south on the Uzboy dry watercourseon the edge of the Karakumdesert (Anon. 1985).
Present range: 1. Khoshyeylag
I; 2. Miandasht
I + Touran V* complex; 3. Bahramgor
IV; 4. Moteh V; 5. Kavir
N’Ajjer II#; 7. Ahaggar
II (Algeria); 8. Possible cheetah tracks seen in the Qattara Depression,
Egypt (Amman
Mts. reserve (proposed:Mali); 10. Ai’r & Tenet-e VIII (Niger); 11. Tibesti Massif (not protected:Chad).

II* complex (Iran); 6. Tassili1993); 9. Adras des Iforas

steppe, a broad zone of bush and grassland where most of
Iran’s cities are located. It snows in the winter. The
Saharan mountains are hyper-arid, but still receive slightly
higher rainfall than the surrounding desert. They are thus
better vegetated and support small permanent waterholes
and antelope populations (Swift 1975, Le Berre 1991).

In India, the Moghul Emperor, Akbar, is reputed to havecollected some 9,000 animals in his lifetime. According toPocock (1939a), the animals were better captured adultfor this purpose, after having learned to hunt from theirmother. By the early 1900s however, Indian cheetahs
had become so scarce that imports of African animals wererequired to sustain the princes’ stables (Divyabhanusinh
1984), as there was no success breeding them in captivity(see also Part II Chapter 5).

Population Status
Global: Category 3(A). Regional: Category l(A). IUCN:
Endangered. Cheetahs were probably extirpated in the following countries during the mid- to late 1900s: Afghanistan, Iraq, Israel, Jordan, Libya, Kuwait, Morocco, Oman,
Pakistan, Saudi Arabia, Syria, Tunisia, Turkmenistan,
Uzbekistan, Western Sahara, and Yemen (Wrogemann1975, Kraus and Marker-Kraus 1991: see Fig. 2). A small,isolated population may persist in Egypt’s Qattara
Depression (IUCN 1976, Kraus and Marker-Kraus
199 1,
Ammann 1993).
De Smet (1989) estimates that “several dozen” cheetahs
persist in the mountains of southeastern Algeria, and it is
not clear whether the population is isolated from that centered on the Air massif 500 km to the south in Niger.
There are no records of cheetahs from the extreme south ofAlgeria (Kowalski and Rzebik-Kowalska
1991, K. de
Smet in Zitt. 1993). Dragesco-Joffe (1993), based on his
travels in the region, estimated the number of cheetahs
remaining in Chad, Mali, and Niger to be between 300 and
500-however,most of these animals are found in the subSaharan dry woodland Sahel region (J. Newby, pers.
comm.). Millington and Anada (199 1) estimated the number of cheetahs in Niger, concentrated in the Air andTermit desert regions and the Sahelian “‘W” National Park,at 200. In Iran, B. Dareshuri estimates the Iranian population to be fewer than 50, with the northeastern provinceof Khorasan being the stronghold (Karami 1992). The
population has declined steeply in recent years: there weresaid to be over 200 cheetahs in Iran in the mid- 1970s (E.
Ferouz, pers. comm. 1974), although some experts consider this figure an over-estimate (P. Joslin, pers. comm.).
Various proposals have been put forward to re-stock
depleted areas with cheetahs of sub-Saharan stock (e.g.

Habitat and Distribution
Cheetahs were once widely distributed across the region,
absent only from extensive sand plains and massifs, andfrom areas of dense tree and shrubby vegetation (Heptnerand Sludskii 1972). At present, only two main population
concentrations can be confirmed: in the southwestern
Sahara and in Iran (Fig. 2). In the southern Sahara, mountain ranges in Algeria, Chad, Mali, and Niger form thecheetah’s stronghold, although they can range far out onto
sandy plains where there is sufficient prey. Cheetahs havebeen observed at elevations up to 2,000 m in the rocky
mountains (Kowalski
and Rzebik-Kowalska
199 1,
Dragesco-Joffe 1993, K. de Smet in Zitt. 1993). In Iran,there are reliable recent records of cheetahs from theprovinces of Khorasan (northeastern part of the country),
Markazi (central), and Fars (southwest) (Karami 1992).
It is possible that cheetahs occur sporadically in otherparts of the Saharan and southwest Asian regions (such as
Egypt’s Qattara Depression, where tracks possibly madeby a cheetah were recently found [Ammann 19931), but
most records date back at least 20 years (see Fig. 2 caption). In southwest Asia, the locations of the greatly
reduced gazelle populations are fairly well-known(East
1992b), and it is unlikely that cheetahs would be overlooked. In north Africa, the situation is more optimistic:
although no longer common, the dorcas gazelle (whichcheetahs in Algeria have been observed to prey upon:
Dragesco-Joffe 1993, K. de Smet in Zitt. 1993) still occurs
widely in certain parts of Egypt, locally in Libya, and in
the southern deserts of Tunisia (East 1992a).
In Iran, cheetahs are found mainly in the central shrub
43

Part I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

LeopardAsia, leopards have so far been studied only in Israel’s
Judean Desert, a pristine mountainous region bordering theDead Sea, where 6-9 individuals have been radio-collaredand monitored since 1979 (Ilani 1990). These leopards
prey mainly on rock hyrax, followed by ibex and porcupine. Ilani (198 1) observed a female leopard hunt hyrax
by leaping blindly over large boulders, surprising a groupof hyrax on her fourth attempt and killing a young male.
Roberts (1977) records an incident of a pair of leopards
attacking a camel in Baluchistan, but describes more typical prey as smaller female and sub-adult Sind ibex andmarkhor, as well as porcupine. Ibex and hyrax were also
reported, along with the Arabian red-legged partridge, tobe the principal prey of leopards in Oman (Daly 1990).
Wild pig were reported as major prey in the forests ofnorthern Algeria (Kobelt 1886, cited in Kowalski andRzebik-Kowalska
1991) and northern Iran (Joslin 1990a).
In the Caucasus mountains, leopards are believed to preyprimarily on wild goats and moufflon (M. Akhverdian inZitt. 1993). In Turkmenistan, the leopard’s range almost
totally coincides with that of Turkmenian sheep (Heptnerand Sludskii 1972), but where these have been depleted
wild boar are the major prey (Lukarevsky 1993).
Leopards from the Arabian peninsula are pale in colorand of small average size (Harrison and Bates 1991).
Further north, in the Judean Desert, one male leopard
weighed 30 kg and two females averaged 23 kg (Ilani
198 1). Leopards attain larger size in the mountains of Iranand central Asia, with recorded weights for males up to
90 kg (Harrington 1977). Leopards in these areas are often
referred to as “snow leopards” in local parlance becauseof their light color and long-haired winter coat (Ognev
1935, Hatt 1959, Harrington 1977)

Israel, India, Turkmenistan, and Uzbekistan), but conserving extant populations is the priority. In addition, reintroduction should not be seriously considered until genetic
comparisons (Hemmer 1988) and environmental impact
evaluations have been carried out. The advice of the IUCN/SSC Reintroduction Specialist Group should be obtained.
Protection Status
CITES Appendix I. National legislation: protected over its
known extant range, and in many historical range states.
Hunting prohibited: Algeria, Egypt, Iran, Kazakhstan,
Morocco,Mali, Niger, Pakistan, Sudan, Tunisia,
Turkmenistan, Uzbekistan. No information: Iraq, Libya,
Mauritania, Jordan, Oman, Saudi Arabia, Syria, Yemen(IUCN Environmental Law Centre 1986, Nichols et al.
1991, E. Mukhina in litt. 1993).
Principal Threats
The cheetahs of Iran and the Sahara exist in very low numbers, divided into widely separated populations. Their low
density makes them particularly vulnerable to reductionof antelope prey through livestock overgrazing and hunting, coupled with direct persecution (cheetahs prey on livestock, especially young camels: K. de Smet, pers. comm.).
While protected areas comprise a key component of cheetah range, management needs to be improved. For example, grazing of domestic stock is reported to be particularly
serious in Iran’s Khosh Yeilagh Reserve (Karami 1992),
once known to hold an important resident cheetah population (Harrington 1977).
Cheetahs native to north Africa and southwest Asia are
not known to be held in captivity.
Action Planning
Projects 36, 37, and 78.

Habitat and Distribution
Leopards are believed to be absent from the true desert ofthe central Arabian peninsula (Harrison and Bates 1991),
although they are found near the Dead Sea, where annual
rainfall is less than 50 mm (Ilani 1990). Pine forest andMediterranean scrub are also suitable habitats for the
species in northwest Africa (Drucker 1990, Kowalski andRzebik-Kowalska
1991), Iran (Joslin 1990a), and theCaucasus (Ognev 1935). Throughout the region they are
confined chiefly to the more remote montane and rugged
foothill areas (Fig. 3), ranging up to 1,800 m in Turkmenistan (Bragin 1990), 3,000 m in Morocco (Drucker 1986),
2,600 m in Saudi Arabia (Biquand 1990) and 3,200 m in
Iran (Misonne 1959).

Leopard, Panthera pardus(Linnaeus, 1758)
Other Names
Panther (English); leopard, panthere (French); Leopard,
Panther (German); leopardo, pantera (Spanish); alym
(Abkhazian); prang, palang, dikho (Afghanistan); nimr
(Arabic);
anzariuts, indz, hovaz (Armenian);
jiki(Georgia); namer (Israel); pling (Kurdish); plang, palang
kouh (Persian); bars (Russian); pars, kaplan, panter(Turkey); koplon (Uzbek).

Population Status
Global: Category 5a(A). Regional: Category 3(A). IUCN:South Arabian subspecies nimr Endangered (Oman, Saudi
Arabia, Yemen); North Persian subspecies saxicolor

Description
and Behavior
See main species account under Sub-Saharan Africa.
Across their wide range in north Africa and southwest

44

Part I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Table 1
Leopard Population Status by Country
Country

Population
Estimate

Reference

Extinct, or No
Resident Populations
A. Serhal in liff. 1993
Hufnagl 1972
Kumertoeve 1975
Shoemaker 1993
M. Reza Khan in litt. 1993

Lebanon
Libya
Syria
Tunisia
United Arab Emirates

Small Populations,
Rare and Threatened
K. de Smet in Mt. 1993
Airumyan and Gasparyan 1976,
M. Akhverdian in Mt. 1993
Alekperov et al. 1977
Osborn and Helmy 1980
Chykovany et al. 1990,
A. Bukhnicashvili in Mt. 1993
H. Mendelssohn
in Mt. 1993
Drucker 1990
Daly 1990
Biquand 1990; S. Biquand, J. Gasperetti,
I. Nader in ht. 1993
Lukarevsky 1990
Akin 1989,199l;
Anon. 1989c,
Ullrich and Riffel 1993; S. Umar in liff. 1993
Lukarevsky 1990
Nader 1989, Biqand 1990

Algeria
Armenia
Azerbaijan
Egypt
Georgia
IsraelMoroccoOman
Saudi Arabia

17

Tajikistan
TurkeyUzbekistan
Yemen

Populations Relatively Larger,
But Still Rare and Confined
to Montane Areas
Afghanistan
IranPakistan
Turkmenistan

Habibi 1977, MacPherson and Fernando
Joslin 1990a
Roberts 1977, Groombridge
1988
Lukarevsky 1990

130~150

No Recent Information
Iraq
KuwaitJordanWestern Sahara

45

1991

Leopard

Part I: Species

Accounts.

Chapter

2. NorthSpecies range: abundance
populations
highly localized.
clProtected
area where
species occurs

Africaand Southwest

reduced,
cl

cl

Asia,

LeopardA

Confirmed

recordA

Unconfirmed

recordProtected
area where
species may occur

Figure 3. Distributionof the leopard(I? pardus)
in north Africa and southwest
Asia.
I. Toubkal V (Morocco);
2. Djurdjura
II; 3. Belezma
II; 4. unconfirmed
observationfrom Ain Sefra (SaharanAtlas) (K. de Smet in litt. 1992);
5. Tassili N’Ajjer II#; 6. Ahaggar
II (Algeria);
7. Air Ten&e VIII (Niger); 8. Gebel Elba IV (Egypt & Sudan); 9. St. Catherine(Moussa)IV(Egypt);
10. Al Fiqrah Protected
Area; Il. Asir V (Saudi Arabia); 12. Judean Desert IV (Israel); 13. Sighting near Alanya in 1991
(Ullrich and Riffel 1993); 14. Leopard shot in 1974 near Beyzpari(Anon. 1989~); 15. Termessos
II (Ullrich and Riffel 1993) (Turkey);
16. Kabardino-Balkarsk
I (Russia); 17. Khosrovsk
I (Armenia);
18. Kiamaky
I; 19. Kavir II” complex;
20. Kolahghazi
I; 21. BakhteganI;
22. Hamoun V; 23. Touran V” complex (Iran); 24. Syunt-Khasardag
I complex;
25. Kopetdag
I; 26. Badhkyz
I (Turkmenistan);
27. Ajar
Valley IV; 28. Pamir-i-Buzurg
IV (Afghanistan).

Indeterminate (Afghanistan, Iran, Turkmenistan).
Other“subspecies”
in the region can also be considered
Endangered: the Anatolian leopard tulliana in western
Turkey, the Caucasus mountains leopard ciscaucasia, andthe Sinai leopardjamisi of southern Israel and the Sinai.
Leopards have fared better than the other big cats-lion,
tiger, cheetah-which
historically occurred in the region.
The tiger is extinct, the lion is represented by a single population in India, and the cheetah’s range is a small fraction of what it once was. However, the future of theleopard is far from secure. Throughout the region, leop-

ards generally exist as small, threatened, and widely separate and isolated populations (Shoemaker 1993).
Protection
StatusCITES Appendix I. National legislation: lacking information. Hunting prohibited: Algeria, Armenia, Egypt,
Georgia, Iran, Israel, Jordan, Morocco, Pakistan, Russia,
Saudi Arabia, Turkmenistan, Uzbekistan. No legal protection: Lebanon, Oman, Tunisia, Turkey, United ArabEmirates. No information: Afghanistan, Azerbaijan, Iraq,
Libya, Kuwait, Syria, Tajikistan, Yemen (IUCN Envir-

46

Part I: Species

onmental Law Centre 1986, Shoemaker 1993; M.
Akhverdian, A. Bukhnicashvili, E. Mukhina, A. Serhal inlitt. 1993).

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Sand

cat

to satisfy its moisture requirements from its prey. Its coatis pale yellow to grey; the tail is ringed and there are darkhorizontal bars on the legs. Sand cats are prolific diggers,
an adaptation not only for hunting fossorial rodents but
for constructing or improving upon the burrows in which
they shelter, such as those dug by the sand fox (M. Abbadi
in Zitt. 1993). Dragesco-Joffe (1993) notes that the sandcat’s claws are not very sharp, as there is little opportunity to sharpen them in the desert, and that impressions of
the claws are often visible in the tracks. The soles of thefeet are covered with a thick layer of wiry black hair (Fig.
4), insulating the foot pads against extremes of heat andcold and allowing easier movement through sand.
Daytime sand surface temperatures in the Sahara during
the summer can reach 52” C (Yunker and Guirgis 1969).
Day air temperatures range up to 58” C in the shade, butnight temperatures are much lower, ranging down to -0.5”
C (Cloudsley-Thompson
1984). In the northern parts ofthe sand cat’s range, it snows in the winter, and temperatures drop as low as -25” C (Heptner and Sludskii 1972).
The sand cat is generally active only at night, according
to the results of a radiotelemetry study in Israel (Abbadi
1992), tracks seen in the central Kara Kum Desert(Bilkevich 1934, cited in Ognev 1935), and activity patterns observed in captivity (Hemmer 1977). Sand cats
have occasionally been observed above ground in day-Principal Threats
Small isolated populations are vulnerable to disruption ofhealthy population dynamics, as has been documented by
Ilani ( 1990) for the leopards of the Judean Desert. In 1978,
the population of roughly 20 individuals-a low number tobegin with-had
a sex ratio of one adult male: 2.5 females.
Since then, four females were killed by humans, and theonly surviving cubs were two males. As of 1989, there had
been no recruitment since 1984, as all cubs born to theone fertile female were killed by the father, and no immigration has been recorded from the adjacent population in
the Negev Desert. Moreover, there were three different
cases of a female mating and producing cubs with her son
and, by 1989, there remained only two adult females in the
population, both too old to breed.
The ungulate prey base throughout the region has inmany places been severely reduced (East 1992a, b), which
probably accounts at least in part for the leopard’s widespread reputation as a killer of domestic stock (Hassinger
1965, Roberts 1977, Harrison and Bates 199 1, Lukarevsky1993). S. Biquand (in litt. 1993) reports predation on
young camels near Medina in Saudi Arabia, and has foundsheep and goat hair in leopard scats. There are numerous
reports of local people going to extraordinary lengths tokill leopards reported in their vicinity, organizing hunting
parties which do not return until the leopard is found andshot (Borner 1977, Habibi 1977, Gasperetti et al. 1986,
Anon. 1989c, Harrison and Bates 199 1, Anon. 1993f).
Action Planning
Projects 38, 39, and 40.

Sand cat, Fe/is margarita
Loche, 1858
Other Names
Chat des sables (French); Sandkatze (German); gato delas arenas, gato de1 Sahara (Spanish); qit el remel, qit
ramli, biss ramli (Arabic); hattul holot (Israel); sevin(Kazakh); peshaya koshka, barchannaya koshka (Russian);qareschtar, aghsheter (Tamahaq: central Sahara); mushuk(Uzbek).
Description
and Behavior (Plate 3)
The sand cat is well adapted to the extremes of a desert
environment and a psammophilic, or sand-dwelling, existence. It lives in areas far from water sources, and is able

Figure 4. The undersideof a sand cat’s paw is protectedfrom
extreme desert temperaturesby a thick covering of fur. The fur
also helps spread the cat’s weight so it can move more easily
over shifting sands (Harrison
1968, Kitchener1991).

47

Part I: Species

Accounts.

Chapter

2. NodhAfricaand Southwest

Asia,

Sand

cat

Weights of wild-caught adults from Turkmenistan
range from 2.1-3.4 kg for males (n=l2) and 1.4-3.1 kg forfemales (n=5) (Heptner 1970). Hemmer et al. (1976) present morphological data which suggest four distinct subspecies: Saharan (margarita), Arabian (harrisoni), central
Asian (thinobia), and Pakistani (schefSeZi). Karyotyping ofa single specimen of each subspecies at Seattle’s
Woodland Park Zoo has yielded preliminary genetic evidence in support of these populations being separate (L.
Werle, pers. comm., cited in Sausman 1991). However,the distribution patterns and habitat requirements of thesand cat are still poorly understood. Hemmer et al. (1976)
note that there could be a number of isolated sub-populations in the Sahara, centered on the various giant discretedune complexes (ergs).

light near their burrows (Lay et al. 1970, Abbadi 1992),
lying on their backs in a posture which, in captivity, isregularly adopted at temperatures above 30” C and presumably helps to shed internal heat. In captivity, sandcats are very sensitive to humidity (Hemmer 1977), and it
is interesting that during six months of radio-tracking, asand cat was only observed resting outside its burrow inthe daytime after several days of rain (Abbadi 1992).
The sand cat’s ears are large and set widely apart andlow on the sides of the head: this trait flattens the sand cat’s
profile hunting in barren areas, and may aid detection of
movements of subterranean prey (Kingdon 1990), as wellas protect the inner ears from wind-blown sand. The tympanic meati (passages from the external ears to the ear
drums: up to 10.5 x 6.8 mm in diameter) and bullae(rounded bony capsules surrounding the middle and internal ears: 2.5-3.4 cm”) are greatly enlarged relative to othersmall felids (Schauenberg 1974). A highly developed
hearing capacity is important for locating prey which, inarid environments, is not only sparsely distributed, but also
found underground.
There are few data on sand cat prey, in part because
their habit of covering their scats with sand (Hemmer
1977) makes them difficult to locate (Abbadi 1992).
Examination of 182 (Sapozhenkov
196 la) and 53(Mambetzhumaev and Palianigazov 1968) stomachs andfeces of sand cats from three central Asian deserts foundthe major prey species to be a diurnal species of the great
gerbil. These gerbils were probably hunted in their burrows at night, which explains the sand cat’s need for keen
hearing. Hearing also plays an important role in intraspecific communication: sand cats make a short, rasping barkin connection with mating activity (Hemmer 1974a,
Abbadi 1992. P. Quillen in litt. 1993). Their diet also
includes birds, reptiles, and arthropods (Heptner andSludskii 1972, Harrison and Bates 1991, Abbadi 1992).
Dragesco-Joffe (1993) says that the sand cat has a reputation amongst Saharan nomads for being a snake hunter,particularly of horned and sand vipers, which they stunwith rapid blows to the head before dispatching with aneck bite. He also notes that sand cats will cover large kills
with sand and return later to feed.
The first radio-telemetry study of the species, whichmonitored four cats for nine months in Israel’s Aravah
Depression (Abbadi 1992), found sand cats to be regularin their behavior. At nightfall, they took up a lookout
position at their den opening, and surveyed the surroundings for about 15 minutes before leaving. They wereactive generally throughout the night, hunting and travelling an average of 5.4 km. Before retiring below groundat dawn, the same lookout position was adopted at themouth of the burrow. Burrows were used interchangeably
by different cats, and the animals did not change burrows
during the day.

Biology
Reproductive season: In the wild, births have beenreported from January-April in the Sahara (DragescoJoffe 1993), in April in Turkmenistan (Ognev 1935,
Heptner and Sludskii 1972) and September-OctoberinPakistan (Roberts 1977), but are not seasonal in captivity(Mellen 1989, Sausman 1991).
Estrus: (C) 5.25 t 0.75 days (n=2).
Estrus cycle: (C) 46 days (n=l) (Mellen 1989).
Gestation: (C) 59-63 days (n=2: Scheffel and Hemmer
1974); 66-67 days (n=2: Mellen 1989).
Litter size: (C) 2.92 k 0.21 (n=25: Mellen 1989); range up
to five (P. Quillen in Zitt. 1993) and possibly eight(Hemmer 1977).
Age at independence: (W) Young sand cats grow rapidly(Heptner and Sludskii 1972), and are thought to become
independent relatively early, perhaps at 6-8 months(Sausman 1991, H. Mendelssohn in litt. 1993).
Age at sexual maturity: (C) 9 (P. Quillen in litt. 1993) to 14
months (Mellen 1989, Green 1991).
Longevity: (C) up to 13 years, but there is a high frequency
of juvenile mortality in captivity (41% of 32 sand cats bornin 1991: Sausman 1991).
Habitat and Distribution
Sand cats are found in both sandy and stony desert(Schauenberg 1974, Hemmer et al. 1976, Gasperetti et al.
1986, Harrison and Bates 199 1, Abbadi 1992, DragescoJoffe 1993). For example, two specimens collected ineastern Egypt came from rather different habitat types.
One was collected on a sandy plain near Lake Nasser withno vegetation in the immediate vicinity; the other was
found in a rocky valley with widely scattered shrubs andtrees (Goodman and Helmy 1986). Heptner and Sludskii

48

Part I: Species

Species

cl

#

range

.
cl

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Sand

cat

Protected area where
species occurs

Protected
area where
species probably
occursFigure 5. Distributionof the sand cat (E margaM@.
1. Tassili N’Ajjer II#; 2. Ahaggar
II (Algeria);
3. Ai’r & Ten&e VIII (Niger); 4. Specimen collected at Armumuit
oasis, Adrar Souttouf Mts.,
Chad (Hemmer
et al. 1976); 5. Djebel Bou-Hedma
II* (Tunisia); 6. Hai Bar Yotvata IV (Israel); 7. Harrat al-Harrah
IV; 8. Tubayq IV;
9. Mahazat
as Sayed I (Saudi Arabia); 10. Wahibah
Sands (proposed:Oman); 11. Specimens
collected from the Al Liwa oasis, Empty
Quarter,
United Arab Emirates (M. Reza Khan in /itf. 1993); 12 Jal az Zhor V (Kuwait);
13. Moteh V (M. Moinian, pers. comm.; cited in
Groves 1990); 14. Touran V* complex (M. Karami, pers. comm.) (Iran); 15. Repetek I*; 16. Krasnovodsk
I (Turkmenistan);
17. Ustyurt I(Kazakhstan);
18. Kyzylkum
I (Uzbekistan);
19. Registan Desert Wildlife Mgt. Reserve (proposed:
Afghanistan).

(1972) describe the sand cat in Turkmenistan as mostabundant amidst extensive sand massifs, as in the central
Karakum where compacted soils are generally absent. The
micro-distribution of the small mammals which form thesand cat’s prey is often clumped around vegetation and,especially during drought years, does not extend onto baresand. However, following rains, the desert blooms andsmall mammals generally expand their ranges (Happold
1984). Sand cats occur only sparsely in the more clayey
desert soils of the Ustyurt and Mangyshlak regions in thenorthern area between the Aral and Caspian Seas (Heptnerand Sludskii 1972).
It is therefore likely that sand cats range throughout the

sandy interior of the Sahara and the deserts of southwest
Asia, but at present there are no specimens from the following countries: Mauritania, Western Sahara, Mali,
Libya, Sudan, Syria, Iraq, Afghanistan, and Iran (although
there is a report from the vicinity of Teheran [Weigel
19611, and two recent reports from the Moteh and Touran
protected areas: Groves 1990, M. Karami, pers. comm.
19921). No ecological explanation for these gaps in sandcat range has been put forward, and they are even more
perplexing on a micro-scale. For example, sand cats are
known from the Hoggar Mountains of southeastern
Algeria (K. Kowalski in litt.; cited in de Smet 1989), theAi’r Mountains of northern Niger (Pocock 1938; J.

49

Part I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Caracallegal protection: Egypt, Mali, Morocco, Oman, Saudi
Arabia, United Arab Emirates. No information: Iraq,
Jordan, Kazakhstan, Libya, Qatar, Tajikistan, Turkmenistan, Uzbekistan, Western Sahara, Yemen (IUCN
Environmental Law Centre 1986), Nichols et al. 199 1,
Belousova 1993; T. Anada, R. Daly, J. Gasperetti, I.
Nader, M. Reza Khan in Zitt. 1993).

Newby, pers. comm. to K. de Smet), and the Tibesti
Mountains of Chad (Hemmer et al. 1976), but not fromthe Adrar des Iforas massif of northeastern Mali (K. de
Smet in Zitt. 1993). There is a record of the sand cat fromthe area between the Hoggar and Air mountains (Hemmer
et al. 1976), so the absence from the Adrar des Iforas issuspect. Similarly, while the sand cat is known from theAravah Depression of southern Israel, it has not beenfound in the Negev Desert sands just to the west (H.
Mendelssohn in Zitt. 1993).
Figure 5 illustrates the probable distribution of the sandcat. The lack of records from Libya and southern
Afghanistan is particularly puzzling (Hufnagl 1972,
Schauenberg 1974, Hemmer et al. 1976), and will probably be proved false with time. In the early 20th century,
confirmed records were available only from northwestern
Africa, so that when the sand cat was found by Ognev(1926) in Turkmenistan, he described it as a new species.
Arabia was the next area presumed to be a major gap in thesand cat’s range, until a living specimen from the Arabianpeninsula was acquired by the London Zoo (Haltenorth
1953, Hemmer 1974a, Hemmer et al. 1976). Finally,
Hemmer et al. (1976) commented on the unusual lack ofrecords for Egypt despite numerous zoological expeditions, but the first specimen was collected in that country
as their article was going to press (Osborn and Helmy
1980, Goodman and Helmy 1986).

Principal Threats
Although the current lack of knowledge about the species’
status and biology makes an assessment premature, thesand cat appears to be one of the least threatened felid
species. Its preferred habitat is not being lost or degraded;
if so-called “desertification”
is a real phenomenon(Stevens 1994), it should actually benefit the species.
Heptner and Sludskii (1972) were of the opinion that sandcat populations in the central Asian deserts were stable andnot threatened, despite harvests at that time of the order of100-200 skins per year. De Smet (1989) reported thatoasis residents in Algeria did not consider it a threat topoultry, and did not trap it to sell as a pet. On the other
hand, Toubou nomads living northwest of Lake Chad consider the sand cat a frequent chicken thief, which readily
enters their camps in the evenings, but they do not generally retaliate due to traditional religious respect for thesmall cats because of their association with the Prophet
Mohammed (Dragesco-Joffe 1993).
Action Planning
Projects 41 and 78.

Population Status
Global: Category 4. Regional: Category 4. IUCN:
Insufficiently Known; schefleli (Pakistan) Endangered.
Although the sand cat has been frequently described as
rare, this may be a result of its harsh environment and nocturnal, subterranean, and secretive habits. For example,
Abbadi (1992) describes the cats’ “freezing” behavior
when disturbed by people, and tendency to close their eyes
against lights at night, making them very difficult to spot.
Despite early reports that the sand cat population ofBaluchistan’s Chagai Desert was devastated by commercial collectors within 10 years after foreign collectors
became aware of its existence (Roberts 1977, Hemmer
1977), more recent information indicates that the sand cat
still occurs widely in the area (P. Paillat, pers. comm. to
S. Biquand 1993).
M. Abbadi (in Zitt. 1993), who carried out the firstradio-telemetry study of the sand cat in Israel, knew of 22
individuals within his 100 km2 study area. The homerange of one adult male was estimated at 16 km2, and overlapped with those of neighboring males (Abbadi 1992).

Caracal, Caracal caracal(Schreber, 1776)
Other Names
Desert lynx (English);caracal (French);caracal,Wiistenluchs (German); caracal, lince africano (Spanish);ajal, anaq al ardh, washag (Arabic); warsal, bousboela,
mousch, nouadhrar, aousak (Berber: Algeria); psk qarh qol(Dari: Afghanistan); hamotro [killer of blackbuck] (Kutchi
dialect of Gujarati: India); siagosh (Persian); karakal(Russian); itfah (Saudi Arabia); orei, ngam ouidenanga(Tamacheq, Toubou [Touareg] : central Sahara); karakulak, step vasagi (Turkish); karakulak (Uzbek).
Description
and Behavior (Plate 3)
Like cheetahs, caracals were trained to hunt for the nobility in India (Sterndale 1884, Sharma and Sankhala 1984).
In general, caracals from this region are somewhat smaller
than those of sub-Saharan Africa, with paler fur in the arid
regions (Harrison and Bates 1991, K. de Smet in Zitt.
1993). Heptner and Sludskii ( 1972) remark that the pelage

Protection Status
CITES Appendix II. National legislation: lacking information.
Hunting prohibited: Algeria, Iran, Israel,
Kazakhstan, Mauritania, Niger, Pakistan, Tunisia. No

50

Part I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

CaracalEstrus: (W) 5-6 days (n=3). Females copulate with several
males in a “pecking order” which is related to the age andsize of the male. One female was found to have matedwith three different males during every estrus period, eachtime the same individuals in the same sequence (Weisbein1989).

of desert caracals bears a surprising resemblance in color
to that of the goitred gazelle. They also note that Turkmen
caracals have tufts of stiff hairs on the paws like the sandcat. Weisbein (in Mendelssohn 1989) also reports thepresence of a dark form in 5- 10% of the caracal populationin central Israel, with adults grey and young kittens almost
black. The average weight of male carcals in Israel is 9.8
2 1.8 kg (n=6); females weigh 6.2 t 0.7 kg (n=5) and aremarkedly smaller than males (Weisbein 1989).
Diet is similar to that reported from sub-Saharan Africa,
consisting mainly of small mammals and birds (Ognev
1935, Roberts 1977, Sharma and Sankhala 1984).
Through scat analysis, prey remains, stomach contents anddirect observation, Weisbein (1989) determined that thediet of caracals in an irrigated agricultural area of Israel
consisted of 62% mammals, 24% birds, 6.1% reptiles, and1.4% insects. In the deserts of Turkmenistan, tolai hareswere the most important prey species (Sapozhenkov 1962,
Ishadov 1983).
Caracals occasionally tackle larger prey, including adultgoitred gazelle (Heptner and Sludskii 1972). Harrison andBates (199 1) note a report from southern Arabia of a caracal killed by a wounded oryx it had attacked. K. de Smet(in Zitt. 1993) found the tracks of a caracal pursuing a dorcas gazelle in Algeria, and caracals to the northwest of Lake
Chad are reputed to hunt these gazelles, hence the local
Toubou name “gazelle cat” (Dragesco-Joffe
1993).
Roberts (1977) notes a record of a caracal stalking a groupof feeding urial in daylight in Pakistan. Caracals have also
been observed to feed on carrion: Mendelssohn (in litt.
1993) describes garbage dumps at poultry farms as richfood sources, and once saw a caracal leap onto a cart ofdead turkeys and select one. A. Livne (pers. comm. cited in
Skinner 1979) observed a caracal chase two sub-adult
striped hyaenas from a donkey carcass.
Weisbein’s (1989) radiotelemetry study in Israel foundthat caracals rested during the day in dense vegetation orrock crevices, and were generally active from dusk todawn and in early morning. Elsewhere, burrows were also
used for shelter (Heptner and Sludskii 1972, Roberts1977). Males travelled an average of 10.4 t 5.2 km (n=40)
per 24-hour period, while females travelled 6.6 t 4.1 km(n=37) (Weisbein 1989). Nocturnal travels up to 20 km
have been documented by following tracks in the Karakumdesert of Turkmenistan (Sapozhenkov 1960).

Age at independence: (W) 9- 10 months (n= 1; Weisbein1989).
Habitat and Distribution
The caracal is widely distributed through the region, absent
only from true desert (Fig. 6). In north Africa, it is common in the humid forest zone of the northern coastal
regions, and is also found in the Saharan mountain ranges(K. de Smet in lift. 1993) and semi-arid woodlands(Dragesco-Joffe (1993). In microhabitat preference, it istypically associated with either well-vegetated or rocky
areas (Heptner and Sludskii 1972, Gasperetti et al. 1986,
Weisbein 1989, A. Johnsingh in Zitt. 1991, Dragesco-Joffe
1993), which provide cover for hunting as well as shelter.
It is often found near water points (Heptner and Sludskii
1972; S. Biquand, H. Mendelssohn in Zitt. 1993), but isapparently capable of satisfying its moisture requirementsfrom its prey (Dragesco-Joffe
1993, J. Gasperetti in Zitt.
1993).
Population Status
Global: Category 5b. Regional: Category 5a(A). IUCN:
Turkmenian caracal Rare. The regional Red Data Booksof the former U.S.S.R. describe the caracal as rare, withthe largest population found in Turkmenistan (estimatedat 250-300 for the country: Belousova 1993). In
Kazakhstan, the northernmost limit of its range, harsh
winters are the limiting factor (Neronov and Bobrov1991). Small populations occur in Uzbekistan along the
Amu-Darya River (Heptner and Sludskii 1972). The
caracal is described as rare in India, the eastern limit of
its range (Pocock 1939a, Sharma and Sankhala 1984, R.S.
Bhadauria in Zitt. 1991). Overall, and especially compared to the larger cats, the caracal is relatively secure,still widespread, and occasionally common.
The only study of a caracal population in the region was
carried out in an agricultural area in Israel’s Negev Desert(Weisbein 1989). Despite a rich prey base supported byirrigation, home ranges were substantially larger than foundin South Africa (where the only other radiotelemetry studies have been carried out). Male home ranges averaged 22 1
* 132 km2 (n=5), and those of females 57 t 55 km” (n=4).
Home range size was positively correlated with bodyweight, and negatively correlated with prey availability.
Male home ranges overlapped substantially (50%), and typically included those of several females. Two dispersals

Biology
Reproductive season: (W) Year-round (Roberts 1977,
Sharma and Sankhala 1984, Weisbein 1989); in the
Sahara, breeding is reported to occur primarily in mid-winter (Jan) (Dragesco-Joffe 1993); in Turkmenistan, kittens
have been found in April-May (Heptner and Sludskii
1972).

51

Part I: Species

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Caracala..
.. ..-

.-

. --

:

. . *a.

..*‘.

l .. . . .

0.

,
-.

Species

range

.
cl

Protected
area where
species occurs*
clProtected
area where
species may occur

Figure 6. Distributionof the caracal (C. caracal)
in north Africa and southwest
Asia.
1. Kiamaky
I; 2. Kavir II* complex; 3. Touran V* complex;
4. Khab-o-Rouchon
I; 5. Hamoun V (Iran); 6. Registan Desert Wildlife
Management
Reserve (proposed:
Afghanistan);
7. Kirthar II complex; 8. Lal Suhanra V*; 9. Cholistan IV (Pakistan);
10. Sari&a
IIcomplex;11. Dhrangadhra(Wild Ass) IV; 12. Ranthambore
II (India); 13. Badkhyz
I; 14. Repetek
I* (Turkmenistan);
15. Ustyurt I(Kazakhstan);
16. Dilek Yarimadisi
II (Turkey);
17. Azraq Desert IV (Jordan);
18. Harrat al-Harrah
IV; 19. Tubayq IV; 20. Asir V (Saudi
Arabia); 21. Jiddat al-Harasis
VI (Oman); 22. Gebel Elba IV (Egypt & Sudan); 23. Zellaf IV; 24. Nefhusa IV (Libya); 25. El Kala V;
26. Chrea II; 27. Djelfa IV; 28. Tassili N’Ajjer II#; 29. Ahaggar
II (Algeria).

were observed: a male migrated 60-90 km south before
establishing a home range, whereas a female remained in
the vicinity of her natal range, with3 her range partly overlapping that of her mother. Twenty caracals, several ofthem transients, were found to utilize an area of 100 km2(with some ranging outside this area), making for a relatively high local density despite the large home ranges.

Tajikistan, Tunisia, Turkey, Turkmenistan, Uzbekistan.
No legal protection: Egypt, Lebanon, Oman, Saudi Arabia,
United Arab Emirates. No information: Afghanistan, Iraq,
Jordan, Kuwait, Libya, Qatar, Syria, Western Sahara(Nichols et al. 1990, IUCN Environmental Law Centre
1986, Belousova 1993; R. Daly, I. Nader, M. Reza Khan,
A. Serhal, S. Umar in Zitt. 1993).

Protection Status
Populations of Asian range states: CITES Appendix I;
African range states CITES Appendix II. National legislation: lacking information. Hunting prohibited: Algeria,
India, Iran, Israel, Kazakhstan, Morocco, Pakistan,

Principal Threats
Caracals prey mainly on small mammals, which are generally not adversely affected by human settlement (LeBerre 1991). However, caracals are capable of taking
small domestic livestock, and surplus killing can result

52Part I: Species

when the animals are attacked in enclosed spaces(Weisbein and Mendelssohn 1990). Such incidents could
lead to vigorous persecution by pastoralists.
Severalauthors have reported caracals to be susceptible to trappingwith fresh bait (Roberts 1977, Gasperetti et al. 1986).
However, Saharan nomadic pastoralists interviewed by
Dragesco-Joffe (1993) stated that problem caracals weredifficult to eliminate because they did not take bait, andmust be chased and treed by hounds. Weisbein (1989)

Accounts.

Chapter

2. NorthAfricaand Southwest

Asia,

Caracalsuggests that caracals are more disposed towards taking
easily acquired prey (e.g. bait, carrion and domestic animals) in the colder months of winter as an energy saving
strategy. His work indicates that, in the absence of heavy
persecution, caracals can adapt well to living in settled
areas in the region.
Action Planning
Project 42.

53

Part I: Species

Accounts.

Chapter

3. TropicalPart I
Species Accounts

Chapter 3
Tropical Asia

Box 1
Vulnerability

Index to Species of the Region (in order of vulnerability)

Species

Habitat Association
St [Mar] (Tot) Score

Tiger, I? Tigris*
Bornean bay cat, C. Bahia*
Clouded leopard, /V. nebulosa *Asiatic golden cat, C. temmincki*
Flat-headed cat, P. planiceps”
Rusty-spotted cat, P. rubiginosus
Fishing cat, P. viverrinus *Marbled cat, P. marmorata”
Leopard, P. pardus
Jungle cat, F. chaus
Leopard cat, P. bengalensis **1:6[3](9)
N: 2 [0] (2)
1:4[4](8)
l:5[3](8)
N: 3 [0] (3)1:7[0](7)
I: 5 [I] (6)N: 3 [I] (4)B: 6 [5] (11)
B: 6 [5] (11)
B: 7 [5] (12)Range(106 km*)
s:
R:
s:
S:
S:
R:
S:
S:
I:
S:
w:

0
-1
0
0
-1
0
0
-1
+I
+I
+I

Score

Geog.

1.99
0.51
2.79
2.66
1.18
0.78
2.33
2.42
4.84
2.69
8.66

-1
-2
-1
-1
-1
-2
-1
-1
0
-1
+I

Body
Size
Score
L
s
M
M
s
s
M
s
L
s
s

-1
+I
0
0
+I
+I
0
+I
-1
+I
+I

Total
Score

Ranking

-2
-2
-1
-1
-1
-1
-1
-1
0
+I
+3

1(A)

1
2(A)
2
2
2
2
2
3(A)
4
5

Key:
* All or most of this species’ range lies within the region

Habitat Association
St = number of strong + significant habitats
N = Narrow (-1); I = Intermediate (0); B = Broad (+I)
[Mar] = number of marginal habitats(Tot) = total number of habitats

Geographic Range (in millions of km2)
R = Restricted(-2); S = Small (-1); M = Medium(0); W = Wide (+I)

Body Size
L = Large (-1); M = Medium(0); S = Small (+I)

(A) = Actively threatened

Regional Criteria
Habitat association:
Narrow = 2-4 habitat types; Intermediate = 6-9 habitat types; Broad = 1 l-l 2 habitat types.
Geographic
range:
Restricted= II million km2; Small = 2-3 million km2; Medium= 4-5 million
Wide = 8-9 million km?
Body size: Large = 35-135 kg; Medium = 7-20 kg; Small = 56.5 kg
See the Introduction

to the Species

Accounts

for explanationof the vulnerability

54

rankingsystem(pp. 2-6).

km2;

Asia

Par? I: Species

Other Names
Tigre (French); tiger (German); tigre (Spanish); lao hu(Chinese); bagh (Hindi, Bengali: India, Bangladesh);rimau, harimau (Indonesia, Malaysia); klaa thorn (Khmer);sua khong, sua lay (Laos); kaduva (Malayalam: India);sher (Persian); tigr (Russian); pedda puli (Telugu, India);seua (Thailand); tag (Tibetan); amba darla (Udege: Amur
River region, Russia).
Description
and Behavior (Plate 7)
Largest of the extant cats and comparable in size to thebiggest of the fossil felids (Mazak 1981), the tiger is also
one of the best-known
large mammals. The reddishorange to yellow-ochre coat with black stripes and whitebelly is immediately recognizable. The tiger is generally
divided into the following subspecies (Mazak 1981):

l

l

l

l

l

l

l

Chapter

3. Tropical

Asia,

Tiger

the tiger in east Asia, from where two major dispersals
took place approximately two million years ago. To thenorthwest, tigers migrated through woodlandsand along
river systemsinto southwestAsia. To the southand southwest, tigers moved through continental southeastAsia,
somecrossingto the Indonesianislands,and othersfinallyreaching India. Herrington (1987) concursthat the South
China tiger may be regardedasa relict population of the“stem” tiger, living in the probable area of origin of thespecies. It has distinctive primitive skull morphology,
including a shortenedcranial region and close-set,moreforward-facing eye sockets.
Stripe patternsdiffer amongindividual tigers and fromone sideof the cat’s body to the other. The stripesvary innumber, as well as width and propensity to split and run
to spots. The dark lines above the eyes tend to be symmetrical, but the markson the sidesof the face can be different. No two tigers have the samemarkings (Sunquist
and Sunquist 1991). Males have a prominent ruff, whichis especiallymarked in the Sumatrantiger.
White tigers have existed in the wild in India. A whitemalecub takenin Rewa,central India, in 1951, wasthe lastrecord. NamedMohan, this tiger becamethe progenitorof
most white tigers now in captivity when bred with adaughter,proving that the albinismis the result of a recessive gene.White tigers have brown stripeson an off-whitebackgroundand ice-blue eyes(Maruska et al. 1987)
Black tigers have been reported occasionally (Burton
1933, Perry 1964, Guggisberg 1975, Mazak 1981), but
the only physicalevidencerestswith a skinrecoveredfrom
illegal traders in Delhi in October 1992, which hasdeep
black on the top of the headand back extending down theflanks to end in stripes(P. Jackson,pers.comm.). It is nottrue melanism,which is found in leopards,jaguars, andmany other cat species,where the entire pelage is black,but may be an expressionof the agouti genewhich causes
mergingof stripes(L. Lyons in Zitt. 1993). Specimenswith
just a few, very broad stripeshave been photographedin
Kanha NP, India (R. Bedi in Zitt.).
The winter andsummerfur of the Amur tiger, aswell asof the extinct tigers of Turkestan and the Caucasus,differs sharply. The hairsin winter grow denseandlong, giving someanimalsa plushor even shaggyappearance.The
winter coat is generally paler, or more ochraceous,than insummer(Heptner and Sludskii 1972).
While tigers are usually solitary, except for femaleswith cubs, they are not anti-social. Males associatewith
femalesfor breedingandhave beenobservedwith females
and cubs when feeding or resting (Schaller 1967,
McDougal 1977, Sankhala 1978, Sunquist 1981, Thapar
1986, 1989). Bragin (1986) quoted reports of tigers
socializing and travelling in groups. A mature male in
Kanha National Park, India, was greetedby a female andcubs andby a sub-adultmale, thought to be from a previ-

Tiger, Panthera tigris(Linnaeus, 1758)l

Accounts.

P. t. tigris (Linnaeus, 1758). Bengal tiger.
Indian subcontinent.
P. t. virgata (Illiger, 1815). Caspiantiger.
Turkey through central andwest Asia.
P. t. altaica (Temminck, 1844). Amur tiger. Amur
River region of Russiaand China, andNorth Korea.
P. t. sondaica(Temminck, 1844). Javan tiger.
Java, Indonesia.
P. t. amoyensis(Hilzheimer, 1905). South China
tiger. South central China.
P. t. balica (Schwarz, 1912). Bali tiger.
Bali, Indonesia.
P. t. sumatraePocock, 1929. Sumatrantiger.
Sumatra,Indonesia.
P. t. corbetti Mazak, 1968. Tndo-Chinese
tiger.
Continental southeastAsia.

Three races-the Caspian (virgata), Bali (balica), andJavan (sondaica) tigers-have becomeextinct sincethe
1950s. Tiger subspecieshave beenevaluated using bothmorphological and molecular methodologies (Hemmer
1978b, 1987; Mazak 1981, 1983; Herrington 1987).
Herrington (1987) was able to distinguish six subspecies
reliably basedon skull measurements(no Caspianor Balitigers were analyzed), although shenoted that there wasconsiderableoverlap of tigris and corbetti, and someoverlap of corbetti and sumatrae. Tiger subspeciesare now
being re-evaluatedusingthe latesttechniquesof molecularanalysis,with samplesbeing collected from wild tigers in
the RussianFar EastandIndia, andfrom captive Sumatranand South China tigers of known origin and bloodline (S.
O’Brien, pers.comm. 1994).
Hemmer (1987) and Mazak (1983) place the origin of55

Part I: Species

Accounts.

Chapter

3. Tropical

Asia,

Tiger

Table 1Size Variation in Tiger Subspecies(Adult Specimens)

(Ma&k

1981)

Subspecies

Weight (kg)Male
Female--* _ _-____---

Total length (m)lFemaleMale_I_____-_------

Skull length (mm)MaleFemale___..
_____-- _-_______I_------

figrisvirgataaltaicasondaica
amoyensis
balicasumatraecorbetti

180-258
170-240
180-306
100-141130-175
90-100100-140150-195

2.7 - 3.1
2.7 - 2.95
2.7 - 3.3
2.48
2.3 - 2.65
2.2 - 2.3
2.2 - 2.55
2.55 - 2.85

329-378316-369341-383306-349318-343295-298295-335319-365

1 MeasurecPbetween

100-160
85-135100-167
75-115100-115650 8075-110100-130

2.4 - 2.65
2.4 - 2.6
2.4 - 2.75
2.2 - 2.4
1.9 - 2.1
2.15 - 2.3
2.3 - 2.55

275-311268-305279-318270-292273-301
263-269263-294279-302

pegs."

aswell asother carnivores,including bears,weighing up to
170 kg, which they have attacked in their winter dens
(Hepter and Sludskii 1972). They readily eat carrion(Schaller 1967).
Tigers usually attack large prey with a stalk from therear, ending with a rush and, sometimes,a spring to bring
down the prey. When seizing and killing prey, the tiger’s
main target is the neck, either the napeor the throat. The
part seizeddependson severalfactors, suchas the size ofthe prey; the size of the tiger; whether the attack is fromfront, rear or side;and the reactive movementsof the prey.
Most observationshave beenof attackson tethered,young
male buffaloes, whose movements are handicapped.
There have beenrelatively few observationsof attackson
free-ranging wild animals. Attack and killing methods
are described by Brander (1923), Champion (1927),
Burton (1933), Corbett (1957), Schaller(1967), McDougal(1977), Thapar (1986), Karanth 1993, Sankhala (1993),
and Seidenstickerand McDougal(l993). Schaller (1967)
noted that adult tigers appearedto be very cautious, andattackedonly when the dangerof injury wasminimal. He
statesthat a tiger characteristically graspsthe throat after
felling its prey, holding on until the animaldies from suffocation. The throat hold protects the tiger from horns,
antlers,andhoovesandpreventsthe prey from regainingits
feet. Sankhala (1993) statesthat tigers prefer to bite theback of the neck, ascloseaspossibleto the skull, killing thevictim by fracturing the vertebrae and compressingthe
spinalchord. Larger animals,however, aregenerallykilledwith a throat bite. For example, Karanth (1993) examined 181 tiger kills and found that most large prey, suchas sambarand gaur, were killed by throat bites. The preyis then usually draggedinto cover, tigers displaying theirous litter. They moved away together (Wright 1989).
However, malesmay kill cubs fathered by other males:
Smith andMcDougal(l99 1) found that the major causeof
death of tiger cubs in Nepal’s Royal Chitwan National
Park wasinfanticide.
Unlike many other cat species, tigers readily enterwater. During hot seasonsthey will lie half-submergedinlakesandpondsduring the heat of the day. In the GangesBrahmaputramangrove delta region of the Sundarbansin
India and Bangladesh,they constantly swim creeks andacrossbroad rivers. Garga (1948) records tigers swimming a 29 km wide river in the Sundarbansand mentions
the possibility that one may have swum 56 km. The
Sundarbanstigers have taken peopleout of boats(Jackson
1991a). Burton ( 1933) records a tiger swimming eightkm from the Malaysian mainland to Penang Island;
Heptner and Sludskii (1972) report the samedistancesswum across the Amu-Darya and Amur rivers in the
Caspianarea and the RussianFar East respectively. In
India’s RanthambhoreTiger Reserve,tigers have charged
into lakes to kill sambardeer, so that both animalswere
momentarily submerged.Crocodileshave beenkilled andeatenby tigers in the area(V. Thapar, pers.comm.).
Tigers hunt mainly between dusk and dawn, but in the
secureconditions of Ranthambhore in the 1980s tigers
frequently huntedduring the day (Thapar 1992). The principal prey acrosstheir rangeconsistsof various speciesof
deer and wild pigs, but U. Karanth (pers. comm.) statesthat in India’s NagarholeNational Park, gaur are the main
prey, including bulls weighing up to 1,000kg. Tigers will
alsoattack young of elephantsandrhinos,and take smaller
species, including monkeys, birds, reptiles, and fish.
Tigers sometimeskill and eat leopardsandtheir own kind,56

Part I: Species

Accounts.

Chapter

3. Tropical

Asia,

Tiger

ern Nepal. The problem is attributed especially to sugar
cane cultivation right to the edge of the park. The canefields provide good cover for tigers, which then come into
contact with agriculturists. Many deaths arise from accidental confrontations in which the tiger makes a defensive attack.
The Sundarbans tigers have had a reputation as maneaters since at least the 17th century (Bernier 1670), butelsewhere man-eating is usually the result of a tiger’s incapacity, through age or injury, to catch normal prey. A
chance encounter in which such a tiger kills someone in a
defensive reaction and feeds on the body may lead it to target people as easy prey. A man-eating tigress may introduce her cubs to human prey. But deaths and injuriescaused by surprised tigers or a tigress defending her cubsfrom intrusion do not usually lead to man-eating. Schaller(1967) agrees with the view of Corbett (1957): “Tigers,
except when wounded or man-eaters, are on the wholevery good tempered. If warnings (growls, rushes, androars) are disregarded, the blame for any injury inflicted
rests entirely with the intruder.” See Part II, Chapter 2 formore discussion of man-eating.

great strength in dragging, even lifting, heavy carcasses.
Pocock (1939a) cites an instance in Burma of a tiger dragging the carcass of a gaur that 13 men could not move.
A tiger eats 18-40 kg of meat at a time (Baikov 1925,
Locke 1954, Schaller 1967) beginning from the rump. If
undisturbed, it returns to its kill for 3-6 days to feed until
little remains (Karanth 1993a). Large prey is taken about
once a week. Sunquist (198 1) estimated frequency ofkilling by females without cubs at once every 8-8.5 days in
Chitwan. Although highly skilled hunters, tigers are often
unsuccessful. They seldom make the effort to press homea failing attack, but Rice (1986) once observed a tiger pursue a wounded sambar for more than two kilometers forjust over two hours in southern India. Schaller (1967)
observed 12 complete stalks, of which only one was successful, and suggested that it was probable that only one
in 20 attacks succeeded. According to V. Thapar (pers.
comm.), one in 10 attacks are successful in Ranthambore,with its high density of prey.
Cooperative hunting has been observed.
Pocock(1939a) said that couples and family groups hunted
together, but gave no references. Thapar (1986) observed
several instances in Ranthambhore. A group of two males
and three females, possibly a family, behaved like lions,taking up positions round a lake where deer congregatedand driving a target animal from one to the other. Corbett(1953) mentions villagers’ reports of two tigers, attacking
in concert, killing a large tusker elephant.
Although lions and leopards also kill humans, tigers
have the greatest reputation as man-eaters, especially in
India. The history of man-eaters (the term is loosely used
to include fatal attacks due to some form of provocation) isreviewed by McDougal(l987).
He quotes average fatalities due to tigers at 85 1 a year between 1902 and 19 10, and1,603 in 1922 alone. The Champawat tiger is said to havekilled 434 people in Nepal and India before it was shot
(Corbett 1952). However, in recent times, with greatly
reduced numbers of tigers, attacks on people have been
relatively rare, except in the Sundarbans mangrove forest
fringing the Bay of Bengal in India and Bangladesh. The
recent annual toll of people in the Indian Sundarbans tiger
reserve has fluctuated between 66 in 1975- 1976, 15 in
1989, and 42 in 1992 (K. Chakrabarty, S.C. Dey, pers.
comm.). Most deaths have been of fisherfolk, wood-cutters, and honey collectors entering the reserve. The high1992 figure is attributed to illegal entry by people, including young children, seeking to benefit from lucrative
prawn harvesting (S.C. Dey, pers. comm. 1992). Earlier,management measures, including the use of human facemasks on the back of the head to deter tigers (which usually attack from the rear) appeared to be reducing the toll
(Rishi 1988, P. Sanyal, pers. comm. 1990).
Since 1978, over 200 people have been killed in thevicinity of India’s Dudhwa National Park, near southwest-

Biology
Reproductive season: (W) Mating takes place year-round,but most frequently from end November to early April(Mazak 198 1); Manchuria: December-February(Ognev
1935, Baikov 1936); India: November-April (Singh 1959,
Sankhala 1967, Schaller 1967, Sankhala 1978). In Nepal,
young born throughout the year in Chitwan NP, with abirth peak from May-July (Smith and McDougal 1991).
Estrus: (C) mean seven days (Sunquist 1981).
Estrus cycle: (W) 15-20 days in Rajasthan, India (Sankhala 1967); this is supported by observations of malefemale association in Chitwan (Smith 1978, Sunquist
1981). (C) 46-52 days (Sadleir 1966); 34-61 days
(Sankhala 1978); 5 1.9 days in Base1 Zoo (Sankhala 1978).
If a litter is lost, estrus occurs within a few weeks (mean 17days, range 10-39, n=3: Sankhala 1978).
Gestation: (C) about 103 days (Sankhala 1978, Sunquist
and Sunquist 1991, Kitchener 1991).
Litter size: (W) mean 2.98 (range 2-5, n=49 in Chitwan
NP: Smith and McDougal 1991); range l-7 (Brander
1923). Observations of females with cubs indicate that 23 is most common (Sankhala 1978). (C) mean 2.9 (n=49litters, range up to 6, in Indian zoos: Sankhala 1978).
Age at independence:
female: Smith 1984).

(W)

18-28 months (male andJuvenile mortality: (W) In Chitwan, Smith and McDougal(1991) found first-year cub mortality to be 34% (n=144
cubs), of which 73% was whole litter loss due to causes

57

Part I: Species

Accounts.

Chapter

3. Tropical

Asia,

Tiger

The geographic distribution of the tiger once extended
across Asia from eastern Turkey to the Sea of Okhotsk(Fig. 1). However, its range has been greatly reduced inrecent times. Currently, tigers survive only in scatteredpopulations from India to Vietnam, and in Sumatra, China,
and the Russian Far East (Fig. 2).

including fire, floods, and infanticide. Mortality in the second year of life was 17% (n=94), of which only 29% waswhole litter loss. Infanticide was overall the most common
cause of cub death.
Age atfirst reproduction: (W) females 3.4 years (n=5),
males 4.8 years, range 3.4-6.8 years (Smith and McDougal
1991); (C) 3-6 years (Sankhala 1967, Schaller 1967).

Population Status
Global: Category 2(A). Regional: Category l(A). IUCN:
Endangered. There may have been 100,000 tigers at theend of the 19th century; a recent mail survey and literature review of the status of the tiger for CITES (Jackson
1993a) concluded that the maximum number is no morethan 7,700. Including “unofficial” institutions such as circuses, there might be more tigers in captivity in the world
now than in the wild.
Of all the range states, India has by far the largest number of tigers. Gee (1964) suggested that it was possiblethat there were 40,000 tigers in India early in this century,compared to about 4,000 by the time he wrote. In 1972,
an official census found positive evidence of fewer than
2,000 tigers in India (Govt. of India 1972), located in fourmain areas of forest: the foot of the Himalayas in north andnortheastern India, the forests of central and eastern India,
and a narrow strip paralleling the southwestern coast. An
intensive conservation program, Project Tiger, was started
shortly thereafter (for more information about Project
Tiger, see Part II, Chapter l), and its 1989 census estimated numbers nationwide at 4,334. However, there has
been widespread poaching in the early 1990s and the mostrecent 1993 census estimates 3,750 tigers (including, aswith the previous total, sub-adults) (Ghosh 1994).
However, it has been officially stated that the population estimates of predators and prey base in India sufferfrom large margins of error (Govt. of India 1993). Karanth(1987, 1993b) is highly critical of the methodology of pugmark identification and counting (see Part II Chapter 3).
Unofficial estimates in 1993 by experienced tiger managers ranged from 2,000 to 4,500 (Jackson 1993a, V.
Thapar, pers. comm.). Including a few hundred tigers in
Nepal (late 1993 estimate 250: C. McDougal, pers.
comm.), Bhutan, Bangladesh, and western Myanmar, thetotal population of Bengal tigers (Pt. tigris) is probably
not more than 4,500 (Jackson 1993a).
Estimates of the number of Indochinese tigers (P. t. corbetti), found from eastern Burma through continental
southeast Asia to Vietnam, range from 1,050 to 1,750(Jackson 1993a) but there are few data. Rabinowitz (1993)
surveyed major protected areas in Thailand between 1987199 1, and estimated the number of tigers in that country
at 250, in sharp contrast to official government estimatesof 450-600 (Anon. 1994~). The Malaysian Wildlife
Department estimated 600-650 tigers in the Peninsula(Anon. 1994~).

Interbirth intervak (W) 20-24 months (n=7: Smith and
McDougal 1991) - 2-2 l/2 years (Sunquist 198 1); but intwo cases when litters were lost in the first two weeks theinterval was eight months (Smith and McDougal 199 1).
Age at last reproduction:
Kleiman 1974).

(C) 14 years (Crandall 1964,

L$etime reproduction: Data collected over nearly 20 years
by the long-term tiger population monitoring project at
Nepal’s Chitwan National Park enabled Smith and
McDougal (199 1) to present pioneering data on lifetime
reproduction in a wild cat species, a critical component of
population viability models. They found the average
reproductive life span of Chitwan tigers to be 6.1 years
for females (n=12; range up to 12.5 years); and just 2.8
years for males (range seven months to six years). For
females, the mean number of offspring surviving to dispersal was estimated at 4.54 (variance 11.48), and the average number of offspring eventually incorporated into thebreeding population was just 2.0 (variance 3.26). For
males, an average of 5.83 of their offspring survived to dispersal (variance 49.97), and 1.99 were incorporated into
the breeding population (variance 6.97).
Longevity: (W) one female was killed in Chitwan when atleast 15.5 years old (McDougal 1991); (C) up to 26 years(Jones 1977).
Habitat and Distribution
The tiger is found in a variety of habitats: from the tropical evergreen and deciduous forests of southern Asia to theconiferous, scrub oak, and birch woodlands of Siberia. It
also thrives in the mangrove swamps of the Sunderbans,the dry thorn forests of northwestern India, and the tall
grass jungles at the foot of Himalayas. Tigers are foundin the Himalayan valleys, and tracks have been recorded in
winter snow at 3,000 metres (Prater 1971). The extinct
Caspian tiger frequented seasonally flooded riverine land
known as tugai, consisting of trees, shrubs, and dense
stands of tall reeds and grass up to six metres in height.
(When hunting in these reed thickets, tigers sometimes
reared up on their hind legs or leaped upward in order to
see their surroundings: Heptner and Sludskii 1972.) The
tiger’s habitat requirements can be summarized as: someform of dense vegetative cover, sufficient large ungulate
prey (Sunquist and Sunquist 1989), and access to water.

58

Part I: Species0qAccounts.

Chapter

3. Tropicaland Sludskii

1972,

Mazgk

Asia,

Tiger

ballcaA

Confirmed

A

Unconfirmed

recordrecordFigure 1. Historical
distributionof the tiger (I? Tigris): mid-l 800s - mid-l
Matjuschkin
et a/. 1980, Ma 1986, Wang and Wang 1986, Lu 1987).

900s(afterHeptner

1979,198l;

Caspiantiger (extinct):
1. Last known tiger in the Caucasus
region killed in 1922 near Tbilisi, Georgia,
after taking domestic livestock(Ognev
1935). 2. Last known tiger in Turkey killed near Uludere,
Hakkari province,in 1970 @stay 1990). 3. Only tiger reported from Iraq
killed near Mosul in 1887 (Kock 1990). 4. Last known tiger in Iran killed in 1959 in Mohammad
Reza Shah (now Golestan)
II (Vuosalo
1976). 5. Tiger killed in 1899 near the Lob Nor basin, Xinjiang, China (Ognev 1935). Tigers disappeared
from the Tarim River basin in
Xinjiang by the 1920s. 6. Tigers disappeared
from the Manas River basin in the Tian Shan mountains,
west of Urumqi, in the 1960s.
7. Last record of the tiger on the Iii River, their last strongholdin the region of Lake Balkhash,dates to 1948. The last record from thelower reaches of the Amu-Darya
river near the Aral Sea was an unconfirmed
observationnear Nukus in 1968 (8) while tigers disappeared
from the river’s lower reaches and the PyzandhValley (9) once a stronghold,in the Turkmen-Uzbek-Afghanborder region by the early
1970s (Heptnerand Sludskii 1972). In the mid-l 800s tigers were killed 180 km northeast of Atbasar, Kazakhstan(10) and near Barnaul,
Russia (11) (Ognev 1935, Mazak 1981: see below for a note on these records).
Java and Bali tigers (extinct):
12. Most of the eight Balitiger specimens
entered the world’s museumsin the 1930s; tigers probably disappeared
from the island by the end of World War II.
Tigers were eliminatedfrom most of Java by the 1940s and were restricted to Meru Betiri II by 1970. Tigers were last observedin Meru
Betiri in 1976 (Seidensticker1986). South China tiger: 13. Tiger killed in Hong Kong in 1942 (Jackson
1991 a); 14. Two young tigers
15. Tiger captured
near Jiexiu, Shanxi province,
Chinakilled near Hangzhou,
Zhejiang province,
China in 1933 (Allen 1938). Amur tiger:(Allen 1938); 16. The tiger probably disappeared
from South Korea during the time of the Korean War (Won 1981); 17. Tiger killed near
Nerchinskiy
Zavod, Russia, in 1884 (Ognev
1935); 18. Tiger observed
near Lake Baikal in 1875 (Guggisberg
1975, Mazak 1981); 19. In
the late 1800s tigers sometimes
crossed the frozen Tatar Strait to Sakhalin Island in the winter (Guggisberg
1975); 20. Northernmost
record (about 60” 40’ N lat.): tiger killed in 1905 on the Aldan River, 80 km north of Ust’ Maya, Russia; fresh tiger tracks were seen in thesame area 5 days later (Ognev 1935, Mazak 1981). These outlying records in the north of the tiger’s range in the former Soviet Union arelocated up to 1,000 km (20) outside the tiger’s permanently
inhabited range, when tigers followed herds of migratory
prey species (reindeer, wild pig). The Kazakhs recognized
this phenomenon
in their region by naming the tiger the “road” or “travelling
leopard” (Heptnerand Sludskii 1972). Bengal tiger: 21. Last known tiger in Pakistan shot in 1906 near Panjnad, Bahawalpur
state (Roberts1977).

59

Part I: Species

Accounts.

Chapter

3. Tropical

Asia,

Tiger

see

UII
FigurePopulations
continuous

generally

Populations
fragmented

generally

2. Present.
cldistribution

Protected
has beenof the tiger.

area whererecorded

species

See text underOccurrencein Protected

Areas.

housed within Chinese zoos (Tilson et al. 1992).
The Amur tiger (P.t. altaica) is virtually confined to theRussian Far East, although a few may survive along
China’s northeast border area, and possibly also in North
Korea (Jackson 1993a: Figs. 2 and 4). The Russian tigers(most located in Primorye territory, with a smaller population in Khabarovsk) have come under increased poaching pressure in recent years as political and economicchange has swept over the region (Anon. 1993a,g-h;
Pikunov 1994). Tigers in Russia in 1994 numbered only
150-200 (A. Amirkhanov,
Deputy Minister for theEnvironment, in Anon. 1994c). A comprehensive census
in the mid- 1980s estimated a minimum of 250 and a maximum of 430 tigers (Pikunov 1988, Bragin and Gaponov1990). The Russian tiger population had fallen as low as
20-30 animals in the 1930s but recovered under protection
from hunting extended in 1947 (Matjuschkin et al. 1980).
There was intense debate in the late 1980s over Russian
proposals to reduce the number of tigers through sport
hunting, with proponents pointing to the increase in tiger

The Amoy, or South China tiger, (P.t. amoyensis), estimated by Lu and Sheng (1986) to number 4,000 in theearly 1950s was virtually extirpated when officiallyhunted as a pest. They state that about 3,000 tigers were
killed in 30 years. Official government statistics showedthat annual average numbers of skins taken dropped from78.6 in the early 1950s to 30.4 in the early 1960s to 3.8
in the early 1970s and to one by 1979, when the government finally banned hunting (Lu and Sheng 1986). Recent
surveys found evidence of tiger presence and reproduction in southern and northern Hunan, northern Guangdong,
and western Fujian. Tiger presence was also noted in eastem Hunan, and was reported recently from central Jiangxi(Koehler 199 1, Gui and Meng 1993). The main areas oftiger distribution are montane sub-tropical evergreen forest
along provincial borders (Fig. 3). The habitat is highly
fragmented, with most blocks less than 500 km”. The total
population size is probably only some 30-80 animals(Jackson 1993a). The captive population is also too small,numbering only around 50 relatively inbred animals, all60

Part I: Species

attacks on livestock and arguing that there were more
tigers than the prey base could support (Jackson 1987,
Pikunov 1988, Bragin and Gaponov 1989, Shchadinov1989). The wave of poaching has ended discussion of thisissue, and the Russian government and NGOs are cooperating to improve anti-poaching protection (Anon. 1993h,
Pikunov 1994).
The Sumatran tiger (Pt. sumatrae) has also sufferedfrom poaching, as well as loss of habitat to human settlement. A Population and Habitat Viability Analysis(PHVA) workshop held in Sumatra estimated the island’s
tiger population at about 400 with relatively goodprospects in five major reserves, and up to 200 scatteredin other areas of the island (Tilson 1992a).
Tigers require adequate prey, cover, and water. Theirranges vary in accordance with prey densities. Whilefemales need ranges suitable for raising cubs, males seek
access to females and have larger ranges. Thus, in areas
rich in prey throughout the year, such as Nepal’s Chitwan
NP and India’s Kanha NP, female ranges of lo-39 km2 andmale ranges of 30- 105 km2 have been recorded (Sunquist
1981), while in the Russian Far East, where prey isunevenly distributed and moves seasonally, ranges are aslarge as 100-400 km2 for females and 800- 1,000 km2 formales (Matjuschkin et al. 1980). Bragin (1986) estimatedadult tiger density at 1.3-8.6 (including young) per 1,000
km2 in the Sikhote-Alin mountains of eastern Russia,
while Karanth’s (1991) review shows that high quality
tropical habitats can support 7- 12 tigers (including young)

Accounts.

Chapter

3. TropicalAsia,

per 100 km2.
The table above shows the range of various density estimates, and indicates the considerable ecological flexibility of the tiger.
Protection Status
CITES Appendix I. The Amur tiger subspecies was
upgraded from Appendix II to Appendix I in 1987.
National legislation: protected over most of its range.
Hunting prohibited: Bangladesh, Bhutan, Cambodia,
China, India, Indonesia, Laos, Malaysia, Myanmar, Nepal,
Russia, Thailand, and Vietnam. No information: North
Korea (Fuller et al. 1991, Jackson 1993a).
Occurrencein Protected Areas
The tiger, like other big cats, probably has little future outside protected areas because of actual and perceived
threats to livestock and human life. Its current range
extends through one of the most densely inhabited regionsof the world, where human populations are rising at an
average of 1.87 percent per annum (WRI/UNEP/UNDP
1992). In India, the human population had increased byover 300 million (nearly 50%) and livestock by over 100
million during the 20 years since Project Tiger began(Govt. of India 1993).
Bangladesh: Tigers are found throughout the Sundarbans
mangrove forests, including the small reserves (total area
320 km2) of Sundarbans East, South & West IV, and may

Table 2
Densities Reported for Tigers in Different Habitats(Adaptedfrom Karanth1991 and Rabinowitz

Tiger

1993)

LOCZWNl

Habitat Type

Ungulate Prey1

Tiger Density*

Nagarhole 11,India
Ranthambore II, lndia
Chitwan II**, Nepal
Kanha 11,lndia
Bengkulu, Sumatra’
Gunung Leuser II*, Sumatra
Huai Kha Khaeng IV, Thailand
Lazovskiy I, Russia4
Sikhote Alin I*, Russia4

Broad-leaved humid forest
Tropical dry forest
Moist monsoon and riparian forest
Moist monsoon forest/meadows
Lowland rain forest
Montane and lowland humid forest
Mixed seasonal forest
Mixed deciduous-coniferous woods
Mixed deciduous-coniferous woods

Very highMedium-high
High
High
Medium-highMedium-low
Low
Low
Very low

I 1.65
IO.00
8.78
6.92

1

Indexed according to Rabinowitz (1993), with prey biomass in Nagarhole(7,658 kg/km*: Karanth 1991) at the high end of the scale.
213 Tiger densities expressed in number of tigers per 100 km? including young and transients.
Those estimates marked 3 include adults only.
4 Data from Matjuschkin ef al. (1980) and Bragin (1986).

61

3.73
1 .I-2.23

ID3
0.6-0.86
0.139‘45

Part I: Species

Accounts.

Chapter

3. Tropical

Asia,

Tiger

MacKinnon 1986: 237-244), but there have otherwise beenno surveys to map tiger distribution in Cambodia (Chim
Somean, Wildlife Protection Office, in Anon. 1994~).

number about 300 (Anisuzzaman Khan in Zitt.) or 460(Farooq Sobhan, Bangladesh Ambassador at Global Tiger
Forum, New Delhi, 1994). They may still occur in Teknaf
VIII, located in the extreme southeastern tip of the country
bordering Myanmar (MacKinnon and MacKinnon 1986).

China: In China, a 1990 survey found South China tiger
signs in 11 reserves (Koehler 1991); a total of 19 fallwithin its present range (Gui and Meng 1993). Total protected area coverage is about 2,500 km2. Gui and Meng
(1993) identify 12 additional sites, with a total area of
6,000 km”, which they recommend for protection (Fig.
3). Sightings of Amur tigers in Changbai Mts. IV* (1,905
km2) in northeastern China were reported in Chinese
newspapers in 1990 (Anon. 1991f; shown in Fig. 2).

Bhutan: Bhutan’s nine lowland protected areas along thesouthern border with India are all believed to contain tigers(Jackson 1993a, Anon. 1994~). Royal Manas II, which
adjoins India’s Manas II**, is the largest and most significant (shown in Fig. 2). Tigers occur at lower elevations in
Jigme Dorji IV, an enormous reserve comprising the entire
northern third of the country (Dorji and Santiapillai 1989).
The Bhutanese government announced a census result of
237 at the 1994 Global Tiger Forum, noting that sometigers are shared with India (Dash0 Penjore Dorji, pers.
comm.). Non-official estimates in 1993 (Jackson 1993a)
put the population at 20-50.

India: India has 21 reserves specifically managed for tigers
which cover over 30,000 km2 and contain about 1,300
tigers, about one-third of the country total of 3,750 (Ghosh1994). Over half this area consists of buffer zones, withhuman settlement, agriculture, and livestock grazing.
Tigers are also found in about 80 other protected areas, inmost of which people and livestock are present. The

Cambodia: Tigers have been recorded in the proposed
Lomphat reserve (shown in Fig. 2; MacKinnon and

270“/ FU JIANJ20GUIZO U

Figure 3. Current
distributionof the South China tiger, with protectedareas andproposed
network(Gui and Meng 1993). 1. Houhe Nature Reserve (20 km*); 2. Mt. Huping IV(440 km*); 3. Wulingyun
Mt. Nature Reserve (108 km*); 4. Badaguang
Mt. IV (180 km*); 5. Dawei
Mt. IV (50 km2); 6. Jinggang
Mts. IV (53 km*); 7. Taoyundong
IV (60 km*); 8. Bamian Mt. IV (42 km*);
9. Chebaling
IV (75 km*); 10. Mang Mt. IV (64 km*); 11. Babao Mt. IV (32 km*); 12. Chenzia IV(78 km*); 13. Dayunyunko
IV (103 km*); 14. Qianjiadong
IV (52 km*); 15. Wuyi Mt IV (53 km*);
16. Wuyi Mts. IV (565 km*); 17. Meihua Mt. IV (221 km*); 18. Mangdan
Mt. Nature Reserve (42 km*);19. Shennongjia(Shewengia)
IV (705 km*); 20. Mt. Fanjing IV (419 km*); 21. Xiaolingzh
IV (21 km*).

62

Part I: Species

Wildlife Institute of India has identified 12 large blocksof remaining forest with the potential to conserve tiger
populations with long-term viability (Johnsingh et al.
199 1: Fig. 5). They contain both state forests, managed fortimber production, and 47 wildlife reserves, including
those specifically managed for tiger. However, one major
reserve, the Melghat Tiger Reserve, one of the first suchreserves specially declared under Project Tiger, is slated tobe reduced by l/3 to just 1,046 km2 in order to accommodate the large number of people living within the reserve(Aziz 1994). Tigers may disappear in a few decades from
56 other reserves because of low numbers and human pressures (Johnsingh et al. 1991). This could mean the loss ofperhaps up to half the 3,000-4,000 tigers currently thought
to survive in India.

Accounts.

Chapter

3. Tropical

Asia,

Tiger

Komsomol'sk-na/

Gavesbidzhan7

Zndunesia (Sumatra): The major reserves for tigers on
Sumatra are Gunung Leuser II* (9,000 km”) in the northwest of the island (Fig. 2), Kerinci Seblat/Barisan Selatan
II complex (along the southeast coast (Fig. 2), Way
Kambas II (shown in Fig. 2), and Berbak IV on the northern coast. Tilson (1992a) notes that habitat within Kerinci
Seblat is significantly fragmented, and tiger populations
are probably also fragmented.

TaPan

Korea, North: Tigers may possibly survive in North
Korea, and Mt. Paekdu IV*, a border area reserve which
adjoins China’s Changbai Mts. IV*, is a likely place.
Figure 4. Distributionof the Amur tiger in the Russian
Far
East (D. Pikunovin litt.). 1. limits of regular records; 2. limits ofnatural habitat; 3. zones of high density; 4. scattered
records.

Laos: Salter (1993) surveyed villages within and near 18
areas which have been proposed as the basis of a national
protected area system. Tigers were reported present near
the majority of villages in all areas.

Russia: Tigers occur, from north to south in Fig. 2, in the
Sikhote Alin* (3,47 1 km”), Lazovskiy (1,165 km”) andKedrovaya Pad (179 km”) I. Unlike most other parts of
its range, the Amur tiger in Russia lives mainly outside
protected areas (Bragin and Gaponov 1989). A survey of
Lazovskiy Reserve in early 1993 estimated the population at 22 tigers (14 adults and eight sub-adults), with perhaps 10 (eight adults and two sub-adults) living on the
periphery (G. Salkina, pers. comm. 1993; Anon. 1993g).
Bragin (1986) estimated the population of the larger
Sikhote Alin Reserve, of which up to l/3 is not suitable interms of vegetation or prey base for tigers, at 25 adults.
Few confine their movements solely to the reserve (Braginand Gaponov 1989). Korkishko and Pikunov (1994) estimated that there were nine tigers (four males [three adult,
one sub-adult]; five females [four adult, one sub-adult])
in the Kedrovaya Pad in 199 1; it is unlikely that they were
all permanent residents. Thus, only about 20% of Russia’s
tiger population is found in protected areas. Outside theseareas, commercial logging and hunting of ungulates are on
the increase.

MaZaysia:
Tigers have been reported from most protectedareas in peninsular Malaysia (Khan 1987). The largest,
Taman Negara II (4,344 kmz), is shown in Fig. 2.
Myanmar: Myanmar’s protected areas have not been surveyed for tiger presence since Salter (1983) reported themas most abundant in Alaungdaw Kathapa II (shown in Fig.
2). Salter (1983) and WCMC (unpubl. data) also mention
tiger presence in other areas, including Shwe-U-Daung,
Shwesettaw and Tamanthi Wildlife Sanctuaries; PidaungGame Sanctuary; Kyaukpandaung, Natma Taung and
Pegu Yomas proposed National Parks; Pakchan proposed
Nature Reserve on the Tenerassim peninsula (shown in
Fig. 2) and Dipayon and Meinmahla Kyun proposed
Wildlife Sanctuaries.
Nepal: In Nepal, tigers are found almost exclusively inRoyal Chitwan II** (shown in Fig. 2), Royal Bardia II, andRoyal Sukhla Phanta and Parsa IV.

63

Part I: Species

Accounts.

Chapter

3. Tropical

Asia,

Tiger

tiger. Maintenance of present habitat is crucial to thetiger’s future, along with protection from illegal killing.
Seidensticker (1986) attributed the extirpation of tigers on
Bali and Java to extensive habitat fragmentation and theinsularization of small habitat blocks and reserves (~500
km2), widespread loss of critical ungulate prey throughdisease, and overhunting by humans.
Tigers are shot or poisoned for livestock predation andfor financial gain. Large numbers of tigers were killed in
the 20th century in Russia and China where they were officially considered pests, and bounties were paid for theirdestruction. In terms of commerce, tigers have traditionally been hunted primarily for their skins: Heptner andSludskii (1972) point to the rising price of tiger skins as
being an important factor leading to their decline in central
Asia and the Russian Far East in the late 1800s to early
1900s. In addition, tiger bone and other body parts areused in traditional Chinese and Korean medicines. In theearly 1900s Russians sold frozen tiger carcasses whole toChinese marketeers and pharmacists (Heptner and Sludskii
1972). Today, the changed political and economic conditions in the former Soviet Union, and what appears to be
a combination of increased demand among Asian consumers coupled with a decreased supply of wild tigers,
have made poaching for bone the pre-eminent threat to
the Amur tiger. Heavy poaching, again primarily for bone,is also taking place in India, and probably elsewhere
throughout the tiger’s range. The tiger bone trade is discussed in detail in Part II, Chapter 4; livestock depredation is covered in Part II, Chapter 2.
Hunting of tigers for sport has also played a role in their
historical decline. Tiger hunting was prevalent throughout
the range from early times. It became very fashionable
when firearms were introduced to the Indian sub-continent, where it was pursued enthusiastically by British officials and Indian upper classes. For example, when King
George V hunted with the Maharajah of Nepal in 19 11, theparty shot 39 tigers in 11 days. The bag record is claimedby the Maharajah of Surguja, who in 1964 wrote to George
Schaller that he had shot 1,150 tigers “only” over his lifetime (Schaller 1967). Russian soldiers moving east in the
19th century hunted tigers as part of their military training
to increase their courage in battle (Heptner and Sludskii
1972). While historical records from India suggest thattiger populations withstood heavy offtakes for long periodsof time (M.K. Ranjitsinh, pers. comm.), tiger populations
became more vulnerable as habitat decreased, particularly
after World War II. Sport hunters from Europe and theAmericas flew into India and Nepal to obtain trophies withlittle official control. Official records in India show that
480 tigers were shot by sport hunters in the years 19661969. It is likely that many more were shot or poisoned.
Hundreds of skins were exported annually before a ban in1968 (Anon. 1994f).

NAMDAPHA-SUNDARBANSNAGARJUNASAGAR

c

,

0

2alkulFigure 5. Proposed
blocks of protectedareas for tigers inIndia (Johnsingh
et a/. 1991). Twelve habitat blocks (including
47 protected areas as well as state forest land) are identified
which have the potential to conserve
large viable tiger populations. The name of one key protectedarea is given for each unit.

Thailand: In Thailand, Rabinowitz (1993) confirmed thepresence of tigers in 22 protected areas, out of 38 visited.
Sixteen reserves were less than 500 km2 in area. He listed
eight forest complexes or sites >2,000 km2 containing
Thailand’s largest tiger populations: Huai Kha KhaengThung Yai IV complex (>12,000 km? shown in Fig. 2);
Nam Nao II complex (>4,000 km2); Kaeng Krachan IVcomplex (>3,000 km2); Thap Lan II complex (>3,000km2); Huai Nam Dang (proposed II) complex (>3,000km2); Khlong Saeng IV complex (>2,000 km2); Mae Tuen
IV complex (>2,000 km2) and Khao Yai II (>2,000 km2).
Vietnam: Evidence for tiger presence has been foundrecently in 14 reserves: Bach Ma Hai Vin and Nam Bai
Cat Tien II; Anh Son, Bu Gia Map, Kon Cha Rang, Mom
Ray, Muong Nhe (Cha), Xuan Nha and Yok Don IV; andMuong Phang, Muong Te, Pong Quang, Pia Oat and Pu
Nhi Reserves (Nguyen Xuan Dang and Pham Trong Anh1992). Muong Nhe, the largest reserve (1,820 km2), isshown on Fig. 2. The others are less than 600 krn2 in area.
Principal Threats
Commercial poaching, a declining prey base due to overhunting, and loss of habitat are the principal threats to the64Part I: Species

On the other hand, subsistence hunting of ungulate preyby local people is now a powerful force driving the tiger’s
decline over large parts of its range. Rabinowitz (1989)
noted an unexpected low abundance of tigers combinedwith a reduced number of banteng, gaur, and sambar in
Huai Kha Khaeng Wildife Sanctuary in Thailand. U.
Karanth (pers. comm.) suggests that, in tropical Asia, it isunlikely that tigers can reproduce successfully at prey densities below 2-5 ungulates per km2 .
Further north, tigers expand their home ranges toaccount for the seasonal movements of a lower density
ungulate prey base. The highest density tiger population in
Russia, in the Lazovskiy Reserve, occurred amidst a relatively high prey density of 2.25 ungulates per km2 (Bragin
1986). However, Amur tigers are naturally vulnerable tosharp declines in ungulate populations during severe winters, and starvation at this time is a common phenomenon.
Hepter and Sludskii (1972) relate reports of emaciatedadult tigers in winter weighing as little as 70 kg: the stomach of one contained nothing but lichens. They report that,in the Primorye region, winters with abundant snow occur
on average once every four years. Such harsh seasonal
conditions increase the precarious situation of the Amurtiger. Since the collapse of the U.S.S.R., poaching of bothtigers and their prey has led to a rapid decline in the population from 250-430 in the mid-1980s (Pikunov 1988,
Bragin and Gaponov 1990) to 150-200 (A. Amirkhanov,
Deputy Minister, in Anon. 1994~)
Severe habitat loss has occurred in this century withthe growth and spread of human populations, settlement,
and activities. Not only have large blocks of tiger habitat
been converted to human use, but wilderness has been fragmented, creating many isolated tiger populations, some sosmall that genetic deterioration is to be feared (Smith and
McDougal 199 1). As Seidensticker ( 1987) declared in his
review of the extinctions of the Bali and Javan tigers, it isdangerous to rely on small, isolated reserves. Large tractsof contiguous habitat are essential to assure the long-term
survival of wild tigers. The problems of conserving tigers
are discussed in Part II, Chapters 1 and 3.

Accounts.

Chapter

3. TropicalAsia,

Borneanbay cat

Description
and Behavior (Plate 9)
The Bomean bay cat is the mystery cat of the family. Its
description rests on just a few skins and skulls, most collected in the late 1880s scattered in several museums
around the world (Sunquist et al. 1994a). Tissue andblood samples for genetic analysis were acquired only inlate 1992, when a female captured by trappers on the
Sarawak-Indonesianborder was brought to the Sarawak
museum on the point of death. The cat weighed 1.95 kg,
but was estimated to have weighed between 3-4 kg whenhealthy (Sunquist et al. 1994b). No observations of thebay cat’s behavior or ecology have been made since Hose(1893).
The Bornean bay cat has two color phases: chestnutred, the more common, and grey (Pocock 1932, Sunquist
et al. 1994b). The coat of the 1992 female was speckledwith black markings (Sunquist et al. 1994a). Her tail was
long: at 391 mm, 73% of head-body length (533 mm). On
all specimens, the backs of the rounded ears are darker-colored, and a whitish stripe runs down the ventral surface ofthe terminal half of the tail. The bay cat resembles the
Asiatic golden cat not only in these characters, but also inskull dimensions, and may well be an island form (Weigel
1961, Hemmer 1978a, Groves 1982). The Asiatic golden
cat occurs widely throughout southeast Asia, including
Sumatra but not Borneo. Borneo has been separated from
Sumatra and other islands on the Sunda Shelf for lO,OOO15,000 years (Sunquist et al. 1994a). Genetic analysis
indicates a close relationship to the Asiatic golden cat(Collier and O’Brien 1985).
Biology
No information.
Habitat and Distribution
Found only on the island of Borneo. Collection and sighting records with fairly precise locations, shown in Fig. 6,
are all from the highlands, and most are near rivers,
although the latter may reflect a collecting bias (Payne etal. 1985; C. Groves, P. Pfeffer, J. Payne in ht. 1993;
Sunquist et al. 1994b). The record from Mt. Kinabalu is an
unconfirmed sighting at 1,800 m (Payne et al. 1985). In
northeastern Kalimantan in the late 1950s P. Pfeffer (inht.
1992) twice saw the fur of the bay cat in Dyak ceremonial caps. S. Yasuma (in ht. 1987, 1988, 1993) has
looked in vain for evidence of the bay cat in the Bukit
Suharto Protection Forest, located 60 km south ofSamarinda in the eastern coastal lowlands of Kalimantan.
According to Hose (1893), dense primary forest is preferred, but recently several biologists have sighted a bay
cat at night in logged dipterocarp forest along the accessroad to the Danum Valley Field Studies Centre in eastern
Sabah (J. Gasis, P. Hurrell, S. Yorath, pers. comm. to J.
Payne 1993).

Action Planning
Projects 12 and 44-56.

Bornean bay cat, Catopuma
badia (Gray, 1874)
Other Names
Chat bai (French); Borneo-katze (German); gato rojo de
Borneo (Spanish); kucing merah (Indonesia, Malaysia);
kucing Kalimantan (Indonesia).

65

Par? I: Species

Accounts.

Chapter

3. Tropical

Asia,

Cloudedleopardseen in 19551957 was visited again in 1986 and 1989.
The forest was still undisturbed and less populated, as mostvillagers had migrated toward the coastal lowlands.
Action Planning
Project 57.

Clouded leopard, Neofelis
nebulosa (Griffith, 1821)

Possible
speciesin the BorneanclA

Confirmed

Figure 6. Possiblecat (C. badia).

range
highlands

.
cl

Danum
(Sabah,Other Names
Panthere longibande, panther-e nebuleuse (French);
Nebelparder (German); pantera longibanda, pantera nebulosa (Spanish);lamchita, gecho bagh (Bengali:
Bangladesh, India); yunbao (Chinese); engkuli (Iban:
Malaysia); machan dahan (Indonesia, Malaysia); shagraw
kai (Kachin: Myanmar); lamchitia (Khas: Nepal); sua one(Laos); thit kyaung, thit-tet kya [tree-top leopard], in kya(Myanmar); rikulau (Rukai, Paiwan: Taiwan); hso awn(Shan); seua laay mek (Thailand).

ValleyVIII
Malaysia)

records

distributionof the Borneanbay

Population Status
Global: Category 2. Regional: Category 1. IUCN:
Insufficiently Known. The bay cat has long been considered rare (Hose 1893). A fauna1 survey of Sabah (Davies
and Payne 1982) found no evidence of the bay cat.
Rabinowitz et al. (1987) interviewed villagers in Sabah
and Sarawak about local occurrence of clouded leopards,
using pictures in a field guide (Payne et al. 1985). While
many informants had seen clouded leopards, leopard cats,flat-headed cats, and marbled cats, none pointed to the picture of the Bornean bay cat (J. Payne in Zitt. 1993). The
trappers who captured the bay cat in 1992 were apparently
aware of its rarity and value to an animal dealer (Sunquist
et al. 1994b).
Protection Status
CITES Appendix II. National legislation: fully protected
over most of its range. Hunting and trade prohibited:Indonesia, Malaysia (Sabah and Sarawak). No legal protection outside reserves: Brunei (Nichols et al. 1991).
Principal Threats
Unknown, probably deforestation (Collins et al. 1991).
On a positive note, P. Pfeffer (in Zitt. 1992) notes that thesame area of eastern Kalimantan where the fur caps were.

66

Description
and Behavior (Plate 8)
The clouded leopard is named after its distinctive markings-ellipses
partially edged in black, with the insides adarker color than the background color of the pelt, andsometimes dotted with small black spots. Pelt color varies
from ochraceous to tawny to silvery grey (Pocock 1939a).
Black and pale, whitish individuals have been reported
from Borneo (Medway
1965, Payne et al. 1985,
Rabinowitz et al. 1987, S. Yasuma in litt. 1993). The
limbs and underbelly are marked with large black ovals,
and the back of its neck is conspicuously marked with two
thick black bars. The tail is thick and plush, encircled withblack rings, and very long, typically equivalent to headbody length (up to 80-90 cm: Pocock 1939a, Legakul andMcNeely 1977, Mehta and Dhewaju 1990). Swinhoe(1862) described the Formosan clouded leopard as a distinct subspecies (FA. brachyurus) on the basis of a shorter
tail length (55-60 cm), but Pocock (1939a) found that taillength is not a consistent criterion. The legs of the clouded
leopard are short, but its canines are relatively the longestof any felid (3.8-4.5 cm: Guggisberg 1975), and have avery sharp posterior edge. Werdelin (1983a) analyzed
morphological characters in the skulls of cats, and concluded that the clouded leopard has attained pantherine
cranial proportions (especially large teeth) without reaching pantherine cranial size. Clouded leopards are intermediate in size between large and small cats: wild adults
have weighed between 1 l-20 kg (Pocock 1939a, Banks
1949, Prater 1971).
The clouded leopard has arboreal talents rivallingPat? I: Species

those of the margay of South America. In captivity, it
has been seen to run down tree trunks headfirst, climb
about on horizontal branches with its back to the ground,and hang upside down from branches by its hind feet
(Hemmer 1968). It probably does some foraging in trees,but mainly uses them for resting (Rabinowitz et al. 1987,
Davies 1990, W. Brockelman in Z&t. 1993). Most photos
taken by camera traps in Sumatra’s Gunung Leuser
National Park, where tigers occur, were at night (Griffiths
1993). There is speculation that the clouded leopard may
be less nocturnal in Borneo, where other large carnivores
are absent (Selous and Banks 1935, Davis 1962,
Rabinowitz et al. 1987). Clouded leopards swim well,and have been found on small islands off Sabah (Davies
and Payne 1982) and Vietnam (Le 1973, C. Santiapillai inlitt. 199 1).
Pocock (1939a) surmises from the clouded leopard’s
long canines and stocky build that it is adapted to take relatively large ungulate prey. Prey has been reported to consist of birds, primates, and small mammals, as well aslarger prey, such as porcupines, deer, and wild boar (Banks
1949, Le 1973, Prater 1971, Rabinowitz et al. 1987, S.
Yasuma in ht. 1993), but the few data collected by scientists have been mainly of primates. In Sabah, a clouded
leopard was observed feeding on a proboscis monkey inthe branches of a small tree in riverine forest (J. Payne in
Zitt. 1992), and one shot in coastal mangrove in northern
Borneo in 1950 had also just killed a large male proboscis
monkey (Davis 1962). Griffiths (1993) found mainly
remains of primates, but also muntjac and argus pheasant,in a small sample of scats attributed to clouded leopards
from Gunung Leuser NP. Clouded leopards have been
observed hunting primates (pig-tailed macaques and gibbons) in Thailand’s Khao Yai National Park (Davies 1990,
W. Brockelman in Zitt. 1993).
Biology
Estrus: (C) average 6 days.
Estrus cycle: (C) average 30 days (n=72).
Gestation: (C) 93 2 6 days (Yamada and Durrant 1989).
Litter size: (C) l-5, most often 3 (n=7 of 9 litters: P.
Andrews in litt. 1993).
Age at first reproduction:
average 26 months.

(C) both males and females

Age at last reproduction: (C) 12 (Yamada and Durrant1989) - 15 years (P. Andrews in Zitt. 1993); most littersborn to males and females between 2-4 years (Yamada andDurrant 1989).
Longevity: (C) average 11, up to 17 years (Prator et al.
1988).

Accounts.

Chapter

3. Tropical

Asia,

Cloudedleopard

Habitat and Distribution
The clouded leopard is usually characterized as being mostclosely associated with primary evergreen tropical rain forest (e.g., Banks 1949, Prater 197 l), but it also makes use ofother types of habitat. Sightings have also been made insecondary and logged forest (Davies and Payne 1982,
Rabinowitz et al. 1987, Santiapillai and Ashby 1988,
Johns 1989, M. Khan in litt. 1991), as well as grasslandand scrub (Santiapillai and Ashby 1988, Dinerstein andMehta 1989). In Burma and Thailand, its presence has
been reported from relatively open, dry tropical forest (C.
Wemmer in Dinerstein and Mehta 1989, Rabinowitz andWalker 199 1). The clouded leopard has been recorded
from mangrove swamps in Borneo (Davis 1962, Davies
and Payne 1982). The clouded leopard has a wide distribution in China, south of the Yangtze (Tan 1984, China
Cat Specialist Group meeting 1992), apparently occurringin a variety of forest types, but there is no information on
habitat preference or ecology across this large portion of its
geographic range (Fig. 7). It has been recorded in the
Himalayan foothills up to 1,450 m (Biswas et al. 1985),
and possibly as high as 3,000 m (Jerdon 1874).
Clouded leopards are remarkably secretive creatures fortheir size. Four animals turned up in different areas of
Nepal in 1989 after more than a century’s hiatus in official observation, having last been recorded in the country
in 1863 (Dinerstein and Mehta 1989). The records extend
the western limit of the range to central Nepal.
Population Status
Global: Category 3(A). Regional: Category 2(A). IUCN:
Vulnerable. Its elusiveness, arboreality, and forest habitat
make the clouded leopard a difficult subject for study (A.
Rabinowitz, pers. comm), and there has been no in-depth
investigation beyond interviews with local residents orforestry workers. In Taiwan, there have been only a handful of sighting reports from hunters since the 1960s; noneof them have been substantiated (Rabinowitz
1988,
Nowell 1991, K.-Y. Lue, pers. comm.). Little forested
habitat remains in Bangladesh and parts of northeastern
India, and numbers are probably low outside protectedareas (Khan 1986, Johnsingh et al. 1991, Choudhury1993). Although it has a wide range in southern China,
suitable forest habitat is generally fragmented in small
patches (J. MacKinnon, pers. comm.).
The status of the clouded leopard is probably healthiest on the island of Borneo (Rabinowitz et al. 1987), possibly because of the absence of tigers and leopards. As
part of a fauna1 survey of Sabah, Davies and Payne (1982)
provided the first (and thus far only) rough estimate of density; they assumed that 12 one-square kilometer study
areas were surveyed adequately so that presence orabsence of clouded leopard would be detected and, on thePart I: Species

Accounts.

Chapter

3. Tropical

Asia,

Cloudedleopard

Nepal, Taiwan, Thailand, Vietnam. Hunting regulated:
Laos. No legal protection outside protected areas: Bhutan.
No information: Cambodia (Nichols et al. 1991; U. Ohn,
R. Salter, C. Santiapillai in litt.).

basis of three records, came up with a density of one individual/4 km?
Protection Status
CITES Appendix I. National legislation: protected overmost of its range. Hunting prohibited: Bangladesh,
Brunei, China, India, Indonesia, Malaysia, Myanmar,(after Santiapillai

Sumatraand Ashby

Principal Threats
Deforestation is the foremost threat, although the serious-

1988)
Species

Areas where there is positive
evidenceof species
presence.
0

Extent of available
positive evidence

range

#
cl

Protected
area where
species may occur

Protected
area where
species occurs

suitable
habitat, but whereof presence
is not available

Figure 7. Distributionof the clouded
leopard(N. nebulosa).
1. Langtang
II (Nepal); 2. Buxa IV(India); 3. Manas IV** (India) + Royal Manas II (Bhutan)
complex; 4. NamdaphaII; 5. Mouling II (India);
6. Rajkandi Forest Reserve;
7. Pablakhali
IV (Bangladesh);
8. Tamathi Wildlife Sanctuary(Myanmar);
9. Nangun River IV; 10. Mt. Fanjing IX*; 11. Wuyi Mts. IV* (China); 12. Yushan II; 13. Tawu Mts. IV(Taiwan);
14. Cat Ba II; 15. Ba Be II; 16. Cut Phuong II; 17. Kon Kai Kinh IV complex;
18. Yok Don IV;
19. Nam Bai Cat Tien II (Vietnam);
20. Lomphat Reserve(proposed:
Cambodia);
21. Xe Bang Nouane(proposed);
22. Muang Son (proposed)(Laos); 23. Nam Nao II complex + Phu Luang IV; 24. Huai
Kha Khaeng IV complex; 25. Khao Yai II complex; 26. Kaeng Krachan IV; 27. Khlong Saeng IV complex (Thailand);
28. Krau IV; 29. Cracker
Range II (Malaysia);
30. Sungai Kayan Sungai MenteranI;
31. Kutai II; 32. Pleihari Martapura
IV; 33. Tanjung Puting II* complex; 34. Gunung Palung I; 35.
GunungPenrisen/Gunung
Niut Game Reserve;
36. Gunung Leuser II*; 37. Torgamba
Production
Forest; 38. Tigapulu
Hills; 39. Kerinci Seblat II; 40. Gumai Pasemah
IV; 41. Barisan Selatan II; 42. Way
Kambas IV (Indonesia).

68Part I: Species

Accounts.

Chapter

3. TropicalAsia, Asiatic

goldencat

ness depends upon further study of the species’ toleranceof various degrees of forest clearance (Rabinowitz et al.
1987). Secondly, the clouded leopard is widely hunted
for its teeth and decorative pelt, and for bones for the traditional Asian medicinal trade. Clouded leopard pelts
were the most commonly available felid pelts in a surveyof black market wildlife traders in southeastern China(Low 1991). Taiwanese were the main buyers. In Taiwan,
where clouded leopards are now either very rare or extinct,
Nowell (1990) reported that small numbers of imported
pelts are sold to aborigines to make traditional ceremonialjackets. Pelts have also been reported on sale in urban
markets from Myanmar, Laos, Vietnam, Cambodia,
Nepal, and Thailand (Salter 1983, Chazee 1990,
Humphrey and Bain 1990, MacKinnon 1990, Van Gruisen
and Sinclair 1992; R. Salter, TRAFFIC Southeast Asia inlitt. 1993). Clouded leopards have been featured on themenu of restaurants in Thailand and China which cater towealthy Asian tourists (Anon. 1988).

streak on the underside.
Very little is known of the golden cat’s behavior andecology. It is predominantly nocturnal (Griffiths 1993);
Pham (1982) most often observed the species in northern
Vietnam between 23-24:00 at night. It is believed to prey
mainly on large rodents, but its diet also includes amphibians and insects (Le 1973), and probably also birds, small
reptiles, and small ungulates such as muntjac and chevrotains. Golden cats have also been reported to prey onlarger animals: the goral in the mountains of Sikkim, India(Biswas and Ghose 1982), wild pig and sambar deer innorth Vietnam (Pham 1982), and young calves of domestic water buffalo (Pocock 1939a, Tun Yin 1967). Griffiths(1993) attributed two scats from Sumatra’s Gunung Leuser
National Park to this species, containing the remains of a
rat and a muntjac.

Action Planning
Projects 58 and 59.

Estrus cycle: (C) 39 days (n=l) (Mellen 1989).

Biology
Estrus: (C) average 6 days (n=2).

Gestation: (C) average 80 days (P. Andrews

in litt. 1993).

Litter size: (C) 1.11 t 0.11 (n=9) (Mellen 1989); range l3 (Guggisberg 1975, Green 1991).

Asiatic golden cat, Catopuma
temmincki (Vigors andHorsfield, 1827)

Age at sexual maturity: (C) 18-24 months-females;
years-males (P. Andrews in Zitt. 1993).

2

Longevity:I (C) up to 20 years (n=12) (Prator et al. 1988).
Habitat and Distribution
Asiatic golden cats are found in tropical and sub-tropicalmoist evergreen and dry deciduous forests, and have occasionally been reported from more open habitats, such as
shrub and grassland (Pham 1982). In the Himalayas, thespecies has been recorded at elevations up to 3,050 m in
Sikkim, India (Biswas and Ghose 1982) (Fig. 8).

Other Names
Temminck’s cat (English); chat dare d’Asie (French);
Asiatische Goldkatze (German); gato dorado asiatico(Spanish); xonali mekoori (Assamese, India); shonali biral(Bengali: Bangladesh, India); jin mao, huang hu, zhi mabao (Chinese); kucing emas (Indonesia); sua meo, sua pa(Laos); kucing tulap, harimau anjing (Malaysia); kya min,kyaung min (Myanmar); hso hpai, miao thon (Shan); suafai [fire tiger] (Thailand).

Population Status
Global: Category 3. Regional: Category 2. IUCN:
Indeterminate. There is little specific information available. The Asiatic golden cat is widely reported as uncommon and threatened by deforestation (Lekagul andMcNeely 1977, Biswas et al. 1985, Khan 1986, R. Salterin Zitt. 1989). Like the clouded leopard, it is foundthroughout much of south-central China, but there havebeen no studies. The largest skin harvests have come from
Jiangxi (234 in 1980-81), Fujian, Hunan, Sichuan, andYunnan (Tan 1984, B. Tan in Zitt. 1991).

Description
and Behavior (Plate 9)
The Asiatic golden cat is remarkably polymorphic in its
pelage. The most common coloration is fox-red to goldbrown, but it can also be black, brown, or grey. There is avariation, thus far reported only from China, of ocelotlike rosettes and spots, which looks so unlike the plain
form that some taxonomists have considered it a separate
species (Weigel 1961, Leyhausen 1979). Pocock (1939a)classified the patterned form as a distinct subspecies ofgolden cat (C. t. tristis) from Sichuan and Tibet, but B. Tan(in ht. 199 1) reports that these forms have been collectedfrom many areas of China. Adults weigh 8.5-15 kg, withmales notably larger than females (Lekagul and McNeely
1977, Tan 1984). The terminal half of the tail has a whitish

Protection
StatusCITES Appendix I. National legislation: fully protected
over most of its range. Hunting prohibited: Bangladesh,
China, India, Indonesia, Malaysia (Penin.), Myanmar,

69

Part I: Species

Accounts.

Chapter

3. TropicalAsia,

Flat-headedlzl

Speciesq

Protected
area
species
occurs.

Protected
species

cat

range
where

area where
probablyoccursFigure 8. Distributionof the Asiatic golden cat (Cm temminck).
1. Gorumara
Wildlife Sanctuary(India); 2. Manas IV** (India) + Royal Manas II (Bhutan) complex; 3. Balphakram
II (India); 4. Rajkandi
Forest Reserve (Bangladesh);
5. NamdaphaII (India); 6. Wolong IV*; 7. Shennongjia
IV*; 8 Wuyi Mts.
IV*; 9. Babao Mt. IV; 10. Nangun River IV (China); 11. Alaungdaw
Kathapa II (Myanmar);
12. Ba Be II;
13. Cut Phuong II; 14. Kong Cha Rang IV + Kon Kai Kinh IV; 15. Yok Don IV; 16. Nam Bai Cat Tien II
complex (Vietnam);
17. Nam Nao II complex;
18. Doi Chiang Dao IV; 19. Salawin IV; 20. Huai Kha
Khaeng IV complex;
21. Khao Luang II (Thailand);
22. Krau IV (Malaysia);
23. Gunung Leuser II*;
24. Way Kambas IV (Sumatra,Indonesia);
25. Nakai Plateau/Nam
Theun (Laos: proposed).

Nepal, Thailand, Vietnam. Hunting regulated: Laos. No
legal protection outside protected areas: Bhutan, Brunei.
No information: Cambodia. (Nichols et al. 1991; U. Ohn,
R. Salter, C. Santiapillai in litt.).

Flat-headed cat, Prionaihrus
planiceps (Vigors andHorsfield, 1827)

Principal Threats
Like the clouded leopard, the golden cat is threatened primarily by deforestation, and secondarily by hunting for
its pelt and bones. Livestock depredation, which usuallyleads to persecution, has also been reported (Prater 197 1,
Lekagul and McNeely 1977).

Other Names
Chat a tete plate (French); Flachkopfkatze (German); gato
cabeciancho (Spanish); kucing hutan, kucing dampak(Indonesia); kucing hutan (Malaysia); gaung bya kyaung(Myanmar); maew pa hua baen (Thailand).

Action Planning
Projects 59 and 60.

Description
and Behavior (Plate 10)
Even more than the fishing cat, the flat-headed cat appears

70

Part I: Species

remarkably adapted to a life of piscivory, or fish-eating(Leyhausen 1979). It has a long, sloping snout and flattened skull roof, and its unusually small ears are set welldown on the sides of the head. It has large, close-set eyes
which provide maximal binocular vision, and the anterior
upper pre-molars are larger and sharper relative to othercats (P2 height and width: P. bengalensis 1.7 & 1.6 mm; P.
planiceps 5.2 and 4.5 mm [Muul and Lim 19701; protocone of P3 also more long and sharp than other cats ofPrionaihus:
Groves 1982). A more developed premolar
is characteristic of mammals that hunt slippery prey, andprovides a better grip (Lyddeker 1896). Finally, the fleshy
sheaths that cover a cat’s claws are shortened in the flatheaded cat, so that only about one-third of each claw iscovered when retracted (Ewer 1973). While the flatheaded cat’s claws do not rub against the ground when
walking, they are always visible. Its toes are more completely webbed than the fishing cat’s (Leyhausen 1979),
and the pads are long and narrow, similar to the Bomean

speciesAccounts.

Chapter

3. TropicalAsia,

Flat-headedcat

bay cat (Pocock 1932b). Muul and Lim (1970), commenting on the cat’s feet and other features, termed it theecological counterpart of a semi-aquatic mustelid, andLeyhausen (1979) has commented on several behavior patterns (prey capture, scent-marking) which are similar to
those of both mustelids and viverrids.
The pelage of the flat-headed cat is thick and soft, andof a reddish-brown color tinged with grey, with the top of
the head more brightly red. Wild adults have weighed 1S2.5 kg (Banks 1949, Muul and Lim 1970). The tail is very
short, only 25-35% of head body length (TL=13-17 cm:
Yasuma and Alikodra 1990).
The stomach contents of an adult shot on a Malaysian
riverbank consisted only of fish (Muul and Lim 1970), andthe stomach of a male killed on a road in a Kalimantan forest reserve contained fish scales and shrimp shells (S.
Yasuma in ht. 1993). In Borneo, flat-headed cats are mostfrequently observed at night along riverbanks, hunting
frogs and fish (Banks 1949; J. Payne, A. Rabinowitz in ht.

is recorded

Protected
area where
species
may occur

Figure 9. Distributionof the flat-headedcat (I? planiceps).
1. Sepilok [Mangrove]
I (Sabah,
Malaysia);
2. Muara Sebuka Nature Reserve (proposed);
3. Kutai II; 4. Bukit Suharto V; 5. Tanjung
Puting II*; 6. Hutan Sambas Nature Reserve(proposed)(Kalimantan,
Indonesia);
7. Similajau
II(Sarawak,
Malaysia);
8. Way Kambas IV; 9. Berbak IV; 10. Kerinci Seblat II complex (Sumatra,Indonesia);
11. Endau Rompin NP (proposed)(Peninsular
Malaysia);
12. Phru Tao Dang Peat
Swamp Forest protectedarea (Thailand: Thai Royal Forest Dept. in litt. 1993).

Pat? I: Species

Accounts.

Chapter

3. TropicalAsia,

Rusty-spottedcat

cat through contamination of its prey. This is a major
problem throughout much of the flat-headed cat’s range(Foster-Turley et al. 1990). In addition, waterways areoften the areas first cleared by people as settlement
expands into the forested areas (Collins et al. 1991).

1993). In captivity, flat-headed cats enjoy a basin of water,
playing or simply sitting in it for hours. They have been
observed to wash objects, raccoon-style. Live fish werereadily taken, with full submergence of the head, and thefish were usually carried at least two meters away, suggesting a feeding strategy to avoid letting aquatic prey
escape back into water (Muul and Lim 1970; M.
Rosenthal, S. Yasuma in litt. 1993).
Although Muul and Lim (1970) reported that their captive cat did not chase after sparrows, the cat at the Lincoln
Park Zoo took live chicks (M. Rosenthal in Zitt. 1993).
Banks (1949) stated that the flat-headed cat was not a poultry raider, but Guggisberg (1975) noted that the only specimen seen by ornithologist B.N. Smythies during his 20
years in Sarawak was shot while chasing chickens. In
addition, M. Khan (in Zitt. 199 1) reports that a female flatheaded cat was captured in Perak, Peninsular Malaysia, in
a trap set to catch cornmon civets preying on poultry.

Action Planning
Projects 61 and 62.

Rusty-spottedcat,Prionailurus rubiginosus(I. Geoff roy Saint-Hilaire,
1831)
Other Names
Chat rougeatre, chat rubigineux (French); Rostkatze(German); gato rubiginosa, gato rojizo (Spanish); bitari
billi (Gujarati: India); kaadu bekku (Kannada: India);pakkan (Malayalam: India); wal balalla, kolla diviya, handun diviya (Sinhalese: Sri Lanka); namali pelli (Tamil:
India); kadu poona, verewa puni (Tamil: Sri Lanka).

Biology
Gestation: (C) approximately 56 days (n=l).
Longevity:1993).

(C) 14+ years (n=2) (M. Rosenthal in litt.

Habitat and Distribution
Most collection records for the flat-headed cat are fromswampy areas, oxbow lakes, and river-me forest (C. Grovesin Zitt. 1993). No research has been done on the species in
the wild; for example, the only information on altitudinalrange for the species (up to 700 m in the Dulit mountainsof Sarawak: Hose 1893) is 100 years old. It may be lessspecialized than presently believed in its habitat requirements, as indicated by sightings in oil palm plantations in
Malaysia, where it apparently hunts rodents (M. Khan inZitt. 1991). It is not known north of the Isthmus of Kra(Lekagul and McNeely 1977, U. Ohn in Zitt. 1993) (Fig. 9).

Description
and Behavior (Plate 10)
The rusty-spotted cat is the cat family’s smallest member.
Males weigh about 1S-l .6 kg, and females 1.1 kg (Phillips
1935, Pocock 1939a). The coat is a short, soft fawn-greywith a rufous tinge, patterned with transverse lines of small
rusty-brown spots which form solid stripes along the backof the head. The tail, which averages about 50% of headbody length, is faintly marked with dark rings (Pocock
1939a).
Very little is known of the rusty-spotted cat’s behavior
in the wild. They are apparently nocturnal (Chakraborty
1978, Pathak 1990, Anon. 199Oc), “lying up during thehours of sunshine in a hollow log, tree or thicket in small
woods of heavy timber or in thick scrub-jungles” (Phillips
1935). They climb well (Stemdale 1884), and in the wild
are frequently
observed in trees (Phillips
1935,
Chakraborty 1978, Anon. 1990~). The diet of the rustyspotted cat has not been properly documented; Phillips(1935) reported without elaboration that it feeds uponsmall mammals and birds. Local people in both Sri Lanka
and India have reported that they are most visible after
heavy rain, when they emerge to feed on rodents and frogs(De Alwis 1973, S. Worah in litt. 1993). They are known
to prey on domestic poultry (Phillips 1935, Pocock 1939a,
J. Zacharias in litt. 1992).

Population Status
Global: Category 2. Regional: Category 1. IUCN:
Insufficiently
Known. The flat-headed cat is seldom
encountered and is believed to be rare.
Protection
StatusCITES Appendix I. National legislation: fully protected
over most of its range. Hunting and trade prohibited:Indonesia, Malaysia, Myanmar, Thailand. Hunting regulated: Singapore. No legal protection: Brunei (Nichols etal. 199 1; U. Ohn in litt. 1993).
Principal Threats
Water pollution, especially by oil, organochlorines, andheavy metals associated with agricultural run-off and logging activities, poses a serious threat to the flat-headed

Biology
Length ofestrus:

72(C) 5 days (n=l).

Part I: Species*.
. -Gestation: 67.6 t 2.0 days (n=4).
:

*

.
*-

Litter size: 1.55 +: 0.25 (n=9) (Mellen 1989).

*

Accounts.

Chapter

3. TropicalAsia,

Rusty-spotted

cat

..

.a*

.:
.

.

.

.
:

I.1.
..

Habitat and Distribution
The rusty-spotted cat is found only in India and Sri Lanka.
Most records are from southern India (Pocock 1939a), butthere are several isolated records from the north of thecountry which are puzzling (Fig. 10). It is difficult to say
whether distribution is continuous throughout India
because the species’ habitat preferences are poorly understood. In Sri Lanka, Phillips (1935) stated that “it is rarely
seen far away from jungles,” while De Alwis (1973) terms
it “the ubiquitous wildcat of Ceylon.. .equally comfortablein the high montane forests of Horton Plains (2,135 m) or
the sizzling sandy wastes of the Hambantota coastline.” In
India, Prater (1971) described its habitat as grassland,
scrub and forest. However, while its presence has been
confirmed in the tropical dry Gir forest (Pathak 1990,
Anon. 199Oc), it appears to be absent from more closed
forest types. According to U. Karanth (in Zitt. 1993), it isprobably not found in the tropical montane rain forest of
the western Ghats. Similarly, residents of 45 villages inthe Dangs semi-evergreen monsoon forest described its
habitat as rocky areas and hill slopes, but not forest edges(Worah 1991).
Perhaps these seeming inconsistencies can be explainedin terms of interspecific competition or ecological separation, although this subject has scarcely been investigated
for the small Tropical Asian cats. The closely related leopard cat is found throughout much of India, but is absent
from Sri Lanka. It is possible that the rusty-spotted cat is
the more common of the two species in the drier, more
open vegetation types of India, while the leopard cat predominates in the moist forests. This would explain theconcentration of rusty-spotted cat records in southern
India, and the infrequent and seemingly isolated reports
from more northern regions. In Sri Lanka, on the other
hand, the leopard cat is absent but the jungle cat occurs,
and is typically found in more open habitats-grass,scrub,and open forest (Phillips 1935).
Rusty-spotted cats show some tolerance of modified
habitat: females with kittens have been found denning ina tea plantation in Sri Lanka (Phillips 1935), and in theattics of houses in southern India surrounded by paddy
fields and coconut plantations (J. Zacharias in litt. 1992).
In the latter case, it was noted that the species was virtually unknown to local residents. A rusty-spotted cat wasphotographed in 1993 in an old farm house in a mango
plantation in Bansda National Park in Gujarat (R. Wirth in
Zitt. 1994). According to Karanth (in Zitt. 1993), rustyspotted cats can be found on farmland throughout southern India’s Deccan Plateau, and on the outskirts of
Bangalore city.

.. .
**. . .
..

q
0A
A

Confwmed

Unconfirmed

,
.

-*
-.*
... ...
*... *-...
*.-..
..
... .
*-..... ..... -j -*.a..... :...;“. . . .;..
*. . . ... .
. . . .. . . ..:-*-.*-.-. ..*..........::

recordrecordFigure 10. Distributionof the rusty-spottedcat (/? rubiginoses).
1. Specimen
collected near Udhampurin 1975(Chakrabot-ty
1978); 2. Gir II complex; 3. Bansda II; 4. Nagarhole
II (India); 5. Wilpattu II; 6. Flood Plains II complex; 7. Sinharaja
IV#; 8. Ruhuna (Yala) II complex (Sri Lanka); 9. Borivali II;
10. Dangs Forest (Worah 1991); 11. Kittens obtained at Seone(Sterndale
1884); 12. Kitten obtained from Tuluka Reserve
Forest, near Purnakat
village, Angul district in 1969 (Wright1984); 13. Rusty-spottedcats living in attics of village houses
near Kochin, Kerala state, India (J. Zacharias
in litt. 1992).

Population Status
Global: Category 3. Regional: Category 2. IUCN:
Insufficiently Known. It has been described as widespreadbut nowhere common (Phillips 1935, Pocock 1939a,
Worah 199 l), as indicated by the patchy and infrequent
nature of collections and observations, but this remains
speculative until basic natural history studies have been
carried out.
Protected Status
Indian population-CITES
Appendix I; Sri Lankan population-CITES
Appendix II. National legislation: fully
protected over its range. Hunting and trade prohibited:
India, Sri Lanka (domestic trade uncontrolled in Sri
Lanka) (Nichols et al. 1991).

73

Part I: Species

Accounts.

Chapter

3. TropicalAsia,

Fishingcat

for fish have been found to be its most frequently taken
prey in Nepal’s Royal Chitwan National Park (D. Smithin Zitt. 1993). Fishing cats are good swimmers, and havebeen observed to dive into water after fish (Breeden 1989),
as well as attempt to scoop them out of water with theirpaws (Leyhausen 1979). Other water-associated prey areprobably taken as well, ranging from crustaceans and mollusts to frogs and snakes. Fishing cats also prey onrodents, small Indian civets, young chital fawns, and wild
pigs (P. Sanyal in litt. 1991, D. Smith in litt. 1993), as wellas domestic goats, calves, dogs, and poultry (Sterndale
1884, Phillips 1935, de Alwis 1973, Bhattacharyya 1988,
Sanyal 1992). Birds are the least frequently taken prey
item in Chitwan (D. Smith in Zitt. 1993). Roberts (1977)
reports that in Pakistan fishing cats have been seen to catchwaterfowl by swimming up to them while fully submerged
and seizing their legs from underneath. A fishing cat wasseen scavenging a cow carcass in India’s Keoladeo
National Park (Haque 1988), and in Chitwan, fishing cats
have been observed to scavenge tiger kills, as well as livestock carcasses (D. Smith, pers. comm.).
Phillips (1935) noted that, in Sri Lanka, fishing cats
could be met “at any hour of the day.”Principal Threats
Deforestation and the spread of cultivation are serious
problems for wildlife in both India and Sri Lanka. As far
as rusty-spotted cats are concerned, it is not known if populations can persist in cultivated landscapes, and individuals which take poultry are vulnerable to persecution (J.
Zacharias in l&t. 1992). A long coat made of rusty-spotted cat fur was found for sale in Kathmandu, Nepal (Van
Gruisen and Sinclair 1992). Early reports on rusty-spotted cats refer to hybridization with domestic cats as a common occurrence (S. Worah in Zitt. 1993), but they havenot been substantiated.
Action Planning
Project 63.

Fishing cat, Prionailurus
viverrinus (Bennett, 1833)
Other Names(French);Fischkatze
Chat pecheur, chat viverrin(German); gato pescador (Spanish); mecho biral, mecho
bagh (Bangladesh); math bagral, bagh dasha (Bengali:
India); bun biral, khupya bagh (Hindi: India); kucing
bakau (Indonesia); sua hay (Laos); kyaung ta nga(Myanmar); math billi (Pakistan); kola diviya, handun
diviya (Sinhalese: Sri Lanka); koddi pulli (Tamil: Sri
Lanka); maew pla (Thailand).

Biology
Reproductive season: (W) in coastal wetlands of northeastern India, peak in mating activity January-February,
with births March-May, but mating also observed in June
(Bhattacharyya 1992).
Gestation: (C) 63 (Ulmer 1968) - 70 days (Mellen 1989).
Litter size: (C) 2.61 k 0.28 (n=13) (Mellen 1989); range
l-4 (Green 1991).

Description
and Behavior (Plate 10)
The fishing cat, with its stocky, powerful build and shortlegs, was given its Latin name on account of its ratherviverrine or civet-like appearance (Bennett 1833). Its pelt
is olive grey, and is patterned with rows of parallel solid
black spots which often form stripes along the spine. Its
tail is very short for a felid, less than half the body length(TL = 23-31 cm, 37% of head-body length (n=5): Pocock
1939a). Females are markedly smaller (6-7 kg) than males(1 I- 12 kg) (Sunquist 199 1). Despite its fishing habits, thefishing cat does not show marked morphological adaptations to capturing or eating fish. ‘Like the flat-headed cat,its claw sheaths are shortened, so that the claws are notcompletely enveloped when retracted. Unlike the flatheaded cat, in which the second upper pre-molar is longand sharp (which enables it to grip slippery prey), the fishing cat, as in most cats, has a much smaller and less-developed tooth. Although webbed feet have often been noted
as a characteristic of the fishing cat, Kitchener (1991)
shows that the webbing beneath the toes is not much moredeveloped than that of a bobcat.
The fishing cat, however, is still appropriately named,

Age at independence: (W) 10 months (Weigel 1975).
Longevity: (C) average 12 years (K. Corbett in Zitt. 1993).
Habitat and Distribution
Fishing cats are strongly associated with wetlands. They
are typically found in swamps and marshy areas, oxbow
lakes, reed beds, tidal creeks, and mangrove areas. They
are more scarce around smaller, fast-moving watercourses(D. Smith, pers. comm.). They have been recorded at elevations up to 1,525 m in the Indian Himalayas (Prater
197 1), where they frequent dense vegetation near rivers
and streams. Although fishing cats are widely distributed through a variety of habitat types (including bothevergreen and tropical dry forest: Rabinowitz and Walker
199 1), their occurrence tends to be highly localized.
The fishing cat also has a discontinuous distribution(Fig. 11). It has long been thought to be absent south of theIsthmus of Kra, but Van Bree and Khan (1992) reported
the capture of a fishing cat in Peninsular Malaysia in 1967;
it died in 1977 in Melaka Zoo. Subsequently, Melisch74Pat? I: Species

cl#Protected
species

area
where
may
occur

A

Confirmed

recordChapter

3. TropicalAsia,

Fishingcatalong the lower reaches of the Indus River, although a few
stragglers penetrate the northeast of the country along theRavi and Sutlej rivers (Roberts 1977).

(1995) drew attention to Swinhoe (1862) in which theauthor mentioned having examined a fishing cat from
Malacca (Melaka). He gave no further details on the specimen, which may possibly have come from elsewhere. He
was mistaken, moreover, in reporting the species’ presence
on Taiwan, a mistake still repeated in the literature over
100 years later (for example, Wang and Wang 1986). Still,
it is possible that the fishing cat is present, but very rare, on
the Malay peninsula. The presence of the species in
Singapore, Borneo, and Bali-forwhich some doubtfulrecords exist (Van Bree and Khan 1992)-deserves further
investigation. There is no record of the fishing cat from
China (Wang and Wang 1986), but it might be found in
Guangxi or Yunnan near the border with Vietnam. In
India, the fishing cat is found in the valleys of the Ganga
and Brahmaputra rivers, and along the upper part of theeast coast and possibly still the southwest coast, but notelsewhere in the peninsula. In Pakistan, it is mainly foundcl

Accounts.

Population Status
Global: Category 2. Regional: Category 2. IUCN:
Insufficiently Known. Fishing cats are locally common
around wetlands. Major systems which potentially support
large numbers of fishing cats include the Sundarbans mangrove forests of Bangladesh and India, the terai region
along the foot of the Himalayas in India and Nepal, thefloodplain of the Ganges and the Brahmaputra,
Cambodia’s Great Lake (Tonle Sap), the coastal floodplains of eastern Sumatra, and the deltas of the Salween,
Irrawaddy, Red, Mekong, and Indus rivers (Sanyal 1983,
Khan 1986; R. Salter, C. Santiapillai, C. McDougal inlitt.). However, all of these areas have been highly
affected by human activities. While fishing cats are report-

Figure 11. Distributionof the fishingcat (I? viverrinus).
1. lndus River delta, Keti Bunder N and S
IV (Pakistan);
2. Dudhwa II; 3. Keoladeo
II**; 4. Bhitar Kanika IV; 5. Sundarbans
I**; 6. JaldaparaIV;
7. Balphakram
II; 8. Pakhui IV; 9. NamdaphaII (India); 10. Sundarbans
E, S, & W IV (Bangladesh);
11. Royal Bardia II; 12. Royal Chitwan II** complex (Nepal); 13. Flood Plains II complex;
14. Maduru
Oya II; 15. Bundala IV; 16. Horton Plains II complex(Sri Lanka); 17. lrrawaddy
river delta (not protected) (Myanmar);
18. Salawin IV; 19. Huai Kha Khaeng IV complex;
20. Phu Hin Rong Kla II;
21. Khao Ang Ru Nai IV; 22. Khao Sam Roi Yot II; 23. Khlong Saeng IV complex; 24. Tarutao Island II(Thailand);
25. Bahau, Negeri Sembilan:
first record for Peninsular
Malaysia(Van Bree and Khan
1992); 26. Way Kambas IV; 27. Berbak IV; 28. Gunung
Leuser II* complex; 29. Kerinci Seblat II complex; 30. Barisan Selatan II complex (Sumatra,Indonesia);
31. Ujung Kulon II (Java, Indonesia);
32.
Tonle Sap [Great Lake] (not protected:
Cambodia);
33. MekongRiver delta (not protected);
34. RedRiver delta (not protected)(Vietnam);
35. Bandipur
II (India) (MacKinnon
and MacKinnon
1986: 240).

75

Part I: Species

Accounts.

Chapter

3. Tropical

Asia,

Marbledcatpeado (Spanish); marbal biral (Bengali: Bangladesh,
India); shih mao, shihban mao, xiao yunbao [small
clouded leopard] (Chinese); kucing batu (Indonesia); kucing dahan (Malaysia); kyaung tha lin (Myanmar); maew
laey hin on (Thailand).

edly common around villages in wetland areas where habitat conversion has not been drastic, such as the outskirtsof Calcutta, where the dominant land use is aquaculture(Sanyal 1992), they do not appear to be so adaptable to rice
paddy and other irrigated forms of cultivation (de Alwis
1973, Dao Van Tien in litt. 1990, K. Mukherjee in litt.
199 1). Along India’s thickly-populatedsouthwestern
coast and in the Indus River basin in Pakistan, fishing cats
are probably on the verge of extinction (U. Karanth, T.
Roberts, B. Wright in litt. 199 1, 1993).
In Java, the fishing cat appears to be restricted to small
numbers in isolated coastal wetlands: there were no records
during recent surveys further inland than 15 km and it must
be considered critically endangered (Melisch et aZ. 1995).
The habitat is threatened by human encroachment for agriculture and aquaculture, and pollution by pesticides.
D. Smith (in Zitt. 1993) recorded home range size forfemales in Nepal’s Chitwan National Park of 4-8 km2(n=3); a single male had a home range of 22 km? Jungle
cats were observed in parts of all four fishing cat homeranges.

Description
and Behavior (Plate 8)
The marbled cat’s coat is very similar to that of the clouded
leopard, although the black-edged blotches on the sides of
its body are less distinct, and black spots on the limbs more
numerous. It also has a long tail, equivalent to or longer
than head-body length (TL = 48-55 cm: Pocock 1939a).
Corbett and Hill (1993) place both species in the genus
Pardofelis, noting that “the unique and complex patternof the pelage is unlikely to be independently derived orprimitive.” Groves (1982) also supports a close relationship, noting that, like the clouded leopard, the marbled
cat’s upper canines are relatively enlarged. However, themarbled cat is less than one-third the weight of the clouded
leopard, has a shorter, rounder skull (Pocock 1932b) andshares an identical karyotype with Lynx, Panthera, andUncia (Wurster-Hilland Centerwall 1982). The evolutionary history of marmorata continues to be a taxonomic
puzzle: Wozencraft’s (1993) revision of the family Felidae
concluded that the classification of the marbled cat should
best be considered incertae sedis, or uncertain.
Very little is known of its behavior, diet, and ecological niche. It is believed to be primarily nocturnal. The few
times marbled cats were observed in the Bukit Suharto
Protection Forest in Kalimantan were in the evening
between 8 and 10 p.m. (Yasuma and Alikodra 1990).
However, in 1994, what is thought to be the first photo ofa marbled cat in the wild was taken during daylight hours
by a photo trap in Thailand’s Huai Kha Khaeng Wildlife
Sanctuary (K. Conforti, pers. comm.). The stomach of a
specimen shot in Sabah contained remains of a small rat(Davis 1962). There was an observation around the turn ofthe century of a marbled cat stalking a bird in a tree
(Guggisberg 1975). Squirrels have been reported in thediet (Ha Dinh Due, Wang Yingxiang, pers. comm.). The
marbled cat has proved to be an adept climber in captivity(Leyhausen 1979).

Protection Status
CITES Appendix II. National legislation: protected overmost of its range. Hunting prohibited: Bangladesh, China,
India, Indonesia, Myanmar, Nepal, Pakistan (Northwest
Frontier), Sri Lanka, Thailand. Hunting regulated: Laos.
No legal protection: Bhutan, Malaysia, Vietnam. No
information: Cambodia (Nichols et al. 1991; U. Ohn, R.
Salter, in litt. 1993).
Principal Threats
Wetland destruction is the primary threat facing thespecies. A survey of the status of Asian wetlands foundthat 50% of over 700 sites were faced with moderate tohigh degrees of threat, including settlement, draining for
agriculture, pollution, and excessive hunting, wood-cutting
and fishing. Severely threatened sites include the estuariesof the Kamataka coast (southwestern India) and the deltasof the Irrawaddy, Indus, Mekong, and Red rivers (Scottand Poole 1989). In addition, clearance of coastal mangroves over the past decade has been rapid in Tropical
Asia (Dugan 1993).

Biology
Gestation: (C) 81 days .

Action Planning
Projects 64 and 65.

Litter size: (C) l-4.
Age at sexual maturity: (C) 21 months (Green 1991).

Marbled cat, Pardofeliis
marntomfa (Martin, 1837)
Other Names
Chat marbre (French); Marmorkatze

Longevity: (C) up to 12 years (Medway

1978).

Habitat and Distribution
The marbled cat is primarily an animal of moist tropicalforest, but there is only anecdotal information on the specificity of its habitat requirements. In Thailand, marbled cats(German); gate jas76

Part I: Species

were not found in dry tropical deciduous forest mosaic
where a study of the carnivore community was carried out,
but were known to be present in adjacent areas of more
extensive mixed deciduous-evergreen forest (Rabinowitz
and Walker 1991). Occurrence in secondary forest has
been noted in Vietnam (Trinh 199 1) and Thailand (B.
Lekagul, pers. comm., cited in Humphrey and Bain 1990),
and Hose (1893) noted that marbled cats were seen frequently in clearings in Sarawak, and were found more
often at low elevations than in the mountains. A marbled
cat was observed in a six-year-old logged forest isolate in
Sabah (Johns 1989). Although most records for the marbled cat in Borneo are from dipterocarp forests, Davies andPayne (1982) observed one individual on a sandy beachstabilized with Casuarina trees and grass, in a remote,swampy mangrove area. Pocock (1939a: 258) reported

Accounts.

Chapter

3. Tropical

Asia,

Marbledcatthat a specimen was captured in a chicken house on the
Barito River, southern Borneo, “in a district which formiles around and for many years had been cleared ofnative forest and planted for rubber and cereals. The animal was living on the river cliff, which consisted of rocks
overgrown with scrub and low bush.”
There are few records on which to base the distribution map (Fig. 12). The sparseness of earlier records isindicated by the following examples: although Pocock(1939a) quotes Horsfield on the marbled cat’s occurrencein hilly regions in Nepal, recent records consist of only asingle specimen circa 1981 from Nawalpur, just to thewest of Royal Chitwan National Park; it has not beenrecorded from the park itself (C. McDougal in Zitt. 1991).
The cat has only recently been recorded from China: aspecimen was collected in China’s Yunnan province in the

Figure 12. Distributionof the marbledcat (/? marmorata).
1. Specimen taken in the Nawal Parasi
district in the early 1980s: only record of species occurrence
for Nepal and westernmost
record of
species distribution(C. McDougal
in litt. 1992); 2. Khangchengdzonga
II (Sikkim, India); 3. Manas IV**
(India) + Royal Manas II (Bhutan) complex;
4. Balphakram
II; 5. NamdaphaII (India); 6. Only record
from China: specimen Shuangjiang,
Yunnan province (Gao eta/. 1987); 7. Ba Be II; 8. Yok Don IV; 9.
Nam Bai Cat Tien II (Vietnam);
10. Phu Luang IV; 11. Khao Yai II complex;
12. Khao Soi Dao IV + Khao
Khitchakut
II; 13. Huai Kha Khaeng IV complex;
14. Khlong Saeng IV complex (Thailand);
15. Kalumba
VIII (Sabah, Malaysia);
16. Gunung
Leuser II* complex;
17. Kerinci Seblat II complex;
18. Barisan
Selatan II complex;
19. Way Kambas IV (Sumatra,Indonesia);
20. Gunung
Penrisen/Gunung
Niut
Game Reserve; 21. Bukit Raya I; 22. Bukit Suharto V (Kalimantan,
Indonesia).

77

Part I: Species

Accounts.

Chapter

3. TropicalAsia,

Leopardjhingfule bagh (Bengali: Bangladesh, India); jin qian bao,
hei bao (melanistic) (Chinese); baghera, tendwa, (Hindi:
India); macan tutul, macan bintang, macan kumbang(melanistic) (Indonesia); khopi (Korean); sua dok, sua dao(Laos); puli (Malayalam: India); harimau bintang, harimau
kumbang (melanistic) (Malaysia); bars (Russian); kotiya(Sinhalese: Sri Lanka); puli (Tamil: India, Sri Lanka).

1970s (Wang and Wang 1986), and B. Tan (in litt. 1991)
writes that there are new reports of its presence in neighboring Guangxi province. Although some distribution
maps have excluded much of southeast Asia (e.g.,
Sunquist 199 1, Corbett 1993) from the marbled cat’s
range, it is present in the lowland forests of southern andcentral Vietnam (Van Peenen 1969; Dao Van Tien, C.
Santiapillai in Zitt. 1991). Husain (1974) thought it
occurred in the Chittagong hill forests of Bangladesh, but
Khan (1986) states that there are no actual records. In
India, the species appears to be restricted to the eastern
Himalayan foothills between 1,500-3,000 m altitude, associated with moist deciduous and semi-evergreen forest
habitats (Biswas and Ghose 1982, Banerjee 1984).

Description
and Behavior (Plate 8)
See main species account under Sub-Saharan Africa.
Melanistic (black) leopards are fairly frequent in populations from the tropical regions in Asia. Although numerous subspecies have been described (Pocock 1930,
Shoemaker 1993) in the region, it is the Amur leopard(P.p. orientalis) which shows the strongest and most consistent divergence in pattern. Leopards from this region(Amur river basin and mountains of northeastern China
and the Korean peninsula) have pale cream-colored coats(particularly in winter), and large (5 x 5 cm on the flanks),
widely spaced (up to 2.5 cm) rosettes with thick, unbroken rings and darkened centers (Pocock 1930, Burger
1970, Wirth 1990b). Leopards from northern China andthe Himalayas also have large rosettes and pale wintercoats, but the rosettes tend to be slightly smaller, moreclosely set, and thinner-edged (Pocock 1930, Dobroruka
1964, 1969; C. Groves, R. Wirth in litt. 1991).
The following weights have been recorded for leopards in the region. Five adult female leopards from China(Sichuan, Anhui and Jilin) weighed an average of 32 kg(Hu and Wang 1984, Gao et al. 1987, Wang 1990); threefemale Amur leopards weighed 25,29 and 43 kg (Gao et
al. 1987). Male Amur leopards have weigh 32-48 kg, withexceptionally large males up to 60-75 kg (Ognev 1935,
Heptner and Sludskii 1972). Females from Sri Lanka
averaged 29 kg (n=7: Pocock 1939a); males from Sri
Lanka averaged 56 kg, with the largest being 77 kg(Phillips 1935, Pocock 1939a). In western Thailand, twomale leopards weighed 60 and 70 kg (Rabinowitz 1989).
Two males from central India weighed 50 and 70 kg(Pocock 1939a).
As elsewhere in their range, the leopards of tropical
Asia have a varied diet (including the occasional young
giant panda: Schaller et al. 1985), but show a preference
for small to medium sized ungulates. Major prey species
include muntjac (Java: Santiapillai and Ramono 1992;
Thailand: Rabinowitz 1989), chital deer (India: Johnsingh
1983; Sri Lanka: Muckenhim and Eisenberg 1973; Nepal
terai: Seidensticker et al. 1990), mountain goats (Pakistan
Himalaya: Schaller 1977), roe and sika deer (Ussuri
region, Russia: Abramov and Pikunov 1974, Korkishko
and Pikunov 1994), hog deer (Nepal terai: Seidensticker
et al. 1990), tufted deer (Wolong, China: Schaller et al.
1985, Johnson et al. 1993b), and langur (south India: U.
Karanth, pers. comm). However, Johnson et al. (1993b)

Population Status
Global: Category 3. Regional: Category 2. IUCN:
Insufficiently Known. The marbled cat may be a naturally rare species. On the other hand, 50 years ago, Pocock(194 1: 476) ascribed the rarity of observation to its forest
habitat and nocturnal habits rather than to real scarcity.
Arboreality can also be added to the factors mitigating
against sightings and collection. A field study is long
overdue.
Protection Status
CITES Appendix I. National legislation: protected overparts of its range. Hunting prohibited: Bangladesh, China(Yunnan only), India, Indonesia, Malaysia, Myanmar,
Nepal, Thailand. Hunting regulated: Laos, Singapore. No
legal protection outside protected areas: Bhutan, Brunei.
No information: Cambodia, Vietnam (Nichols et al. 199 1;
U. Ohn, R. Salter in Zitt.; China Cat Specialist Group meeting 1992).
Principal Threats
The degree of forest clearance the species can tolerateneeds investigation (see Part II Chapter 1). The marbled
cat is probably opportunistically hunted, but specimens are
not commonly observed in local wildlife markets (Low
1991, TRAFFIC Southeast Asia in Zitt. 1993).
Action Planning
Projects 59 and 66.

Leopard, Panthera pardus(Linnaeus, 1758)
Other Names
Panther (English); leopard, panthere (French); Leopard,
Panther (German); leopardo, pantera (Spanish); chita bagh,78

Pat-t I: Species

Accounts.

Chapter

3. TropicalAsia,

Leopardbeen observed to breed in June-July (summer) and givebirth in September-October(Shibnev 1989). In Sri
Lanka, breeding is believed to take place during the dry
season (May-July)(Santiapillai et al. 1982). In the
Himalayas, mating may take place mainly in winter, as
rasping calls are heard more often at this time (A.
Johnsingh in litt. 1993).

found a shift over a seven year period in Wolong fromtufted deer to bamboo rats, although they were not certain
whether the shift reflected individual prey preferences ofdifferent leopards, a decrease in deer, an increase in bamboo rats, or an increase in the vulnerability of the rats due
to a bamboo die-off.
Where tigers are present, the much smaller leopards
tend to be few (Schaller 1967, 1972; M.K. Ranjitsinh, pers.
comm.). This is not a strict rule; Korkishko and Pikunov(1994) concluded that an increase in the number of tigers
in Russia’s Kedrovaya Pad Reserve did not affect the leopard population. In Nepal’s Chitwan National Park, leopards and tigers coexist by hunting at different times and for
different prey, as well as by utilizing different vegetation
complexes (Seidensticker
1976). The leopard takes
smaller prey (generally less than 75 kg: Seidensticker
1976, Johnsingh 1983), in a manner similar to the foodresource partitioning found for lions and leopards in the
Serengeti (Bertram 1982) and the Gir Forest (R. Chellamin Zitt. 1993). Leopards are more tolerant than tigers oftemperature extremes and dry environments (Santiapillai
and Ramono 1992)-for example, they are more common
in seasonally dry tropical monsoon forest than tigers,which are dependent upon permanent water sources(Kleiman and Eisenberg 1973, Sunquist 1981, Johnsingh
1983, Rabinowitz 1989).
Rabinowitz (1989) found a relatively high proportion ofprimate remains (12%) in 237 leopard scats analyzed fromThailand’s Huai Kha Khaeng Wildlife Sanctuary. Big cats
prefer not to hunt primates when alternative, more accessible prey species are available and abundant (Seidensticker 1983), and the relatively high rate of predation
found by Rabinowitz may be due to competition withtigers for muntjac. In the dry deciduous forest of Huai Kha
Khaeng, the canopy is relatively open and primates may
necessarily have to do more travelling on the ground(Rabinowitz 1989). Pocock ( 1939a) describes leopards
catching langur monkeys by feigning a move to climb atree, leading them to jump to the ground to escape, where
they could be more easily caught. Observation of the way
langur troops leap in all directions between ground andtrees when alarmed suggests that the monkeys may havedeveloped a technique to confuse the leopard (P. Jackson,pers. comm. 1993).
Average daily movement for a radio-collared adultmale in Thailand was 2 km, and he was active 66% of theday. There was no strong trend towards either nocturnal or
diurnal activity (Rabinowitz 1989). Leopards tend to be
more nocturnal in proximity to human settlement (A.
Johnsingh in litt. 1993).

Age at independence: (W) 12-l 8 months (Eisenberg and
Lockhart 1972, Muckenhim and Eisenberg 1973, Sunquist
1983, Pikunov and Korkishko 1989).
Age atfirst reproduction: (W) 2-3 years (Sunquist 1983,
Pikunov and Korkishko 1989).
Interbirth interval: (W) average 20-21 months (n=6: Royal
Chitwan NP).
Mortality: (W) Seidensticker et al. (1990) found high juvenile mortality among leopards living at the edge of Royal
Chitwan NP: mean litter size when cubs were up to onethird of adult size was 2.3 (n=3), but for cubs at one-half to
two-thirds adult size it was 1.3 (n=6).
Habitat and Distribution
The region encompasses a broad spectrum of environments, and leopards occur in most of them (Fig. 13).
Leopards are found throughout the Indian sub-continent
with the exception of deserts, the Sundarbans mangroves,and densely settled areas (Khan 1986, Johnsingh et al.
1991)-although
they occur on the outskirts of Bombay
adjoining Borivli National Park (P. Jackson, pers. comm.).
Leopards range throughout most of China as well (Tan
1984, Wang and Wang 1986, Gao et al. 1987, Ma Yiqing
in Zitt. 1993). In the Himalayas, leopards are sympatricwith snow leopards up to 5,200 m (Jackson 1984)
although they more commonly live below the tree line
(Roberts 1977, Green 1987).
In Indonesia, leopards are found only on Java and thetiny offshore island of Kangean; fossil remains date toabout one million years ago (Hemmer and Schutt 1973).
Van Helvoort et al. (1985) suspect that the leopard wasintroduced to Kangean Island, which is situated further
from Java than Bali, where leopards do not occur.
Seidensticker (1986) speculates that leopards (and tigers)
are probably absent from the island of Borneo due the lackof a large ungulate prey base, and that leopards were“squeezed out” from the islands of Bali by the presenceof tigers, and from Sumatra by an abundance of otherfelids (seven species).
For a big cat, the leopard is remarkably persistent in
the face of human settlement, especially considering thehigh human population densities found throughout muchof this region. P. Jackson (pers. comm. 1995) saw a leopard on the outskirts of Pakistan’s capital, Islamabad. They
are still found (in low numbers) throughout Java-despite

Biology
Reproductive season: (W) Seasonal throughout the region
except the tropics (Prater 1971). Amur leopards have79Pat-t I: Species

Accounts.

Species

Chapterrange: abundance

3. TropicalAsia,

Leopardreduced,Figure 13. Distributionof the leopard(I? pardus)
in tropicaland east Asia. 1. Margalla Hills V (Pakistan);
2. Kishtwar II;
3. Corbett II; 4. Kumbhalgarh
IV; 5. Gir II complex;
6. AnamalaiIV; 7. Nagarjunasagar
Srisailam IV; 8. Melghat IV; 9. Sanjay II;
10. NamdaphaII (India); 11. Manas IV** (India) + Royal Manas II (Bhutan) complex;
12. Pablakhali
IV (Bangladesh);
13. Wilpattu II (SriLanka); 14. Alaungdaw
Kathapa II (Myanmar);
15. Huai Kha Khaeng IV complex;
16. Khlong Saeng IV complex (Thailand);17. Taman
Negara II (Peninsular
Malaysia);
18. Ujung Kulon II; 19. Meru Betiri II (Java); 20. Pulao Kangean Game Reserve (Kangean
island,Indonesia);
21. Xe Bang Nouane (proposed);
22. Phou Khao Khouay (proposed:
Laos); 23. Nam Bai Cat Tien II; 24. Ba Be II (Vietnam);
25. Royal Chitwan II** complex; 26. Shey-Phoksundo
II complex (Nepal); 27. Medog IV; 28. Gaoligong
Mt. IV; 29. Xishuangbanna
IV;
30. Wolong IV*; 31. Baishui River I; 32. Shennongjia
IV*; 33. Mt. Fanjing IX*; 34. Chebaling
IV; 35. Wuyi Mts. IV*; 36. Mt Jiulong IV;
37. Guniujiang
IV; 38. Mazongling
IV; 39. Liupan Mt IV; 40. Lipan Mt. IV; 41. Luya Mt. IV; 42. Wuling Mt. IV; 43. Changbai
Mts. IV* (China);
44. Kedrovaya
Pad I (Russia); 45. Mt. Paekdu IV* (North Korea); 46 Mt. Chiri V (South Korea).

the fact that the island is one of the most densely populatedareas in the world.

hunting. Leopards are now confined to one isolated habitat block in Bangladesh (Khan 1986), and have been
greatly reduced in the mountains of northern Pakistan(Roberts 1977). Santiapillai et al. (1982) estimated thenumber of leopards in Sri Lanka at 400-600 based on densities of one adult per 20 (Clark 1901) to 30 km*(Eisenberg and Lockhart 1972) in remaining forest habitat.
They believe that numbers have fallen by 75% since theturn of the century.
The Russian range of the Amur leopard, Pp. orientalis,shrank dramatically between 1970- 1983, as leopards disappeared from the southern parts of the Sikhote-Alinmountains, a stronghold of the tiger, losing about 80% ofPopulation Status
Global: Category 5a(A). Regional: Category 3(A). IUCN:
Amur leopard subspecies orient&s and Sri Lankan subspecies kotiya Endangered; North Chinese leopard japonensis Vulnerable; Javan leopard melas Indeterminate.
Seidensticker (1986) suggests that leopards have increased
throughout the region as tigers have declined. Leopards
are better able to survive outside protected areas, but inmost cases populations can be expected to show a declining trend due to habitat loss, depletion of prey, and direct80

Pat? I: Species

Chapter

3. TropicalAsia,

Leopardlast record of an Amur leopard is from 1969, when a leopard was captured on the slopes of Odo Mountain, in SouthKyongsang province. Tracks have been seen on the Chii
and Sorak Mountains, indicating that they have not completely disappeared (Won 1988, Won Pyong-Oh in litt.
1993). In North Korea, Amur leopards may still survive in
the northern mountains (Won 1968, Prynn 1980, Won
Pyong-Oh in Zitt. 1993). Pikunov and Korkishko (1994)
consider poaching to be the main factor currently limiting
leopard numbers in Russia, and the Amur leopard must
now be considered critically endangered.
Based on density estimates of one leopard per 10 km2
in moderately suitable habitats and one per 5 km2 in favorable habitats, Santiapillai and Ramono (1992) estimatedthe Javan population to be 350-700 animals. Its strongholds are in the protected areas shown in Fig. 13, as well as
in more remote montane areas. These densities are considerably higher than Rabinowitz’s (1989) estimate of one
per 25 km2 in tropical dry forest in Thailand; however, thetheir former range (Pikunov and Korkishko 1989). Justprior to their disappearance, Heptner and Sludskii (1972)
had stated that the range of the Amur leopard hadremained relatively stable over the past century. Anothercensus was conducted in 199 1, which documented a minor
loss of range in the south since 1983 (Korkishko
andPikunov 1994: compare Figs. 14 and 15). Overall, thepopulation has remained relatively stable over the last
decade, at a very low level. Korkishko
and Pikunov(1994) estimate numbers at no more than 28-3 1 (6-9 males[4-6 adults, 2-3 sub-adults]; 19 females [14 adults, fivesub-adults]; three sub-adults of unknown sex: Korkishko
and Pikunov 1994).
Amur leopards are now believed to be practicallyextinct in the mountainous regions of China’s northernmost province, Heilongjiang (Ma Yiqing in Zitt. 1993),
although some may persist in the Changbai Mountains in
Jilin province along the North Korean border (D. Prynn,
pers. comm. in Shoemaker 1993). In South Korea, the/AnvuiZ

Accounts.

R.

/‘SuchanFigure 14. Distributionof the Amur leopard
in Russia in 1971. S.P. Kucherenko
in Heptner and Sludskii(1972: 221). 1. Boundaryof region of permanent
residence;
2. Best sites of habitation;
3. Region of very rare
and temporary
intrusions beyond zone of permanent
occurrence;
4. Directions of individual intrusions from
China during 1930 to 1970.

81

Part I: Species

Accounts.

Chapter

3. TropicalAsia,

Leopardleopard may occur at higher densities in Java since thetiger’s extinction in the mid- 1900s (Hoogewerf 1970).
The ability of leopards to persist despite habitat loss andfragmentation is best illustrated by the case of India, where
leopards have been estimated to number some 14,000,
with half in protected areas (Wildlife Institute of India
unpubl. data). Between 1982-1989, 170 people were
killed by leopards, with the majority in the Kumaon andGarhwal hills of Uttar Pradesh, where Corbett (1948)
hunted man-eating leopards in the early part of this century(Johnsingh et al. 1991). Leopards are also common in thefoothills of the Nepalese Himalayas, despite a dense
human population.
Rabinowitz (1989) found that male leopards in the Huai
Kha Khaeng WS maintained slightly overlapping homeranges of 27-37 km*, while females had ranges of 1 l-17
km* within the ranges of males. One male’s home range
was largest (17- 18 km*) during the early rainy season
months of June and July, and smallest (4.4 km*) during theheavy rains and floods of September and October. In
Nepal’s Royal Chitwan NP, Seidensticker et al. (1990)
found similar-sized female home ranges of between 7- 13
km* . Pikunov and Korkishko (1989) reported that homeranges of Amur leopards, based on snow tracking, havebeen estimated at 50-300 km*.

China

Figure 15. Distributionof the Amur leopard
in Russia in
1991 (Korkishko
and Pikunov1994). The city of Vladivostok,shown in Figure 14, is located about 75 km southeastof the cityof Ussuriisk, just off the edge of this map. 1. border of Kedrovaya
Pad I; 2. border of Barsovy Special Reserve;
3. border of a proposed national park on the Shufan plateau; 4. location of leopard
tracks in 1991.

Protection Status
CITES Appendix I. National legislation: fully protected
over most of its range. Hunting prohibited: Bangladesh,
China, India, Indonesia, Laos, Malaysia, Pakistan, Russia,
Sri Lanka, Thailand. Hunting regulated: Nepal (a low
level of trophy hunting was permitted until recently: C.
McDougal in iitt. 1992). No legal protection outside protected areas: Bhutan. No information: Cambodia, North
Korea, South Korea, Myanmar, Vietnam (Nichols et al.
1990, Shoemaker 1993, R. Salter in ht. 1993).

traditional medicine, is widespread in the region (Sayer
1983, Anon. 1986, Ma 1986, Barnes 1989, Anon. 1990b,
Chazee 1990, Humphrey and Bain 1990, Low 199 1, Anon.
1992g, Santiapillai and Ramono 1992, Johnson et al.
1993b, Korkishko and Pikunov 1994). While habitat loss
is still a significant threat, the leopard does well in secondary growth, and is not as vulnerable as other felids toforest clearance (Johns 1989).
The Amur leopards of Russia are additionally threatened by the small size of the population: father-daughterand sibling matings have been observed on two occasions(Korkishko 1986). Korkishko and Pikunov (1994) foundthat the average litter size (measured by tracks in snow)
fell from 1.75 in the winter of 1973, to 1.6- 1.75 in the winter of 1983, and to 1.O in the winter of 199 1. They pointout that it cannot be determined at present whether thedrop is due to genetic factors, such as a decline in fertility,
or is merely a demographic fluctuation.

Principal Threats
Leopards are threatened by a depleted wild ungulate preybase in many areas- such as the Russian Far East
(Korkishko and Pikunov 1994)-and are persecuted when
they turn to livestock. Domestic stock has been found to
be a major component of leopard diet outside protectedareas (Schaller 1977, Seidensticker et al. 1990). Seidensticker et al. (1990) studied leopards living at the sharplydemarcated boundary of Chitwan NP, and concluded thatthe availability of domestic livestock allowed leopards tolive at a higher density than could be supported by wild
prey. However, high juvenile and adult mortality, coupled with suitable habitat left unoccupied for extended
periods after a resident’s death, indicated that the leopard
population was having a difficult time maintaining its
numbers due to persecution.
Illegal commercial hunting, for pelts and for bones for

Action Planning
Projects 67 and 68.

82

Part I: Species

Accounts.

Chapter

3. TropicalAsia,

Junglecat

thickets lined with old cane leaves and fur.
Jungle cats are frequently observed in the daytime.
They feed primarily on rodents (Allayarov 1964, Schaller
1967, Heptner and Sludskii 1972, Roberts 1977, Khan and
Beg 1986, A. Johnsingh in Zitt. 1991), including large
rodents such as the introduced coypu (weight 6-7 kg) in
Eurasia (Dal 1954). Heptner and Sludskii (1972) note that
200 cats were caught in traps over a period of 14 years in
the vicinity of a coypu fur-farming operation. Jungle catsalso take hares, birds, reptiles, amphibians, insects, and the
young of larger mammals such as chital or wild pig(Rathore and Thapar 1984). They are strong swimmers,
and will dive to catch fish (Mendelssohn 1989), or toescape when chased by man or dog (Heptner and Sludskii
1972). One cat in India, observed hiding in a bush while
stalking a group of grey jungle fowl, appeared to makedeliberate clockwise movements of its head, rustlingleaves and attracting the curiousity of the birds (Tehsin andTehsin 1990).

Jungle cat, Fe/k chaus
Schreber, 1777
Other Names
Swamp cat, reed cat (English); chat des marais, chat dejungle (French); Rohrkatze, Sumpfluchs (German); gato
de la jungla, gato de 10s pantanos (Spanish).
Tropical Asia: jongli mekoori (Assamese, India); wab, banberal (Bengali: Bangladesh, India); conglin mao, limao(Chinese); sembalado [a cat living on the boundary of avillage] (Gujarati: India); jangli billi, ban bilao, khattas(Hindi: India); kaadu bekku, bokana kotti (Kannada:
India); meo pa (Laos); kattu poocha (Malayalam: India);baoga (Marathi:
India); kyaung ba, taw kyaung(Myanmar); wal ballala, handun diviya (Sinhalese: Sri
Lanka); kadu poona (Tamil: India, Sri Lanka); maew pa,
sewa kratay (Thailand).

Biology
Reproductive season: (W) Mating behavior reported in
Ott in southwestern India (A. Johnsingh in Zitt. 199 1);
January-February in central Asia (Allayarov 1964). Births
reported in early May in Armenia (M. Akhverdian in Zitt.
1993.)

North Africa and Southwest Asia: bizoon el berr, qatwahshee (Arabic); ehegna katu (Armenia); chel pshigi(Azerbaijan); smuncha (Dari: Afghanistan); gurbeh siah,
gurbeh i kuhi (Farsi: Iran); lelianis cata (Georgian); pishik(Iraq); kamish mishiki (Kazakh); bizoon, pesheela-kaywee, pisheek-kaywee, kitkakive, kithakaywee (Kurdish);kamish suloosunu (Kyrgyz);
kameshovy
kot, haus(Russian); saz kedisi (Turkey); sabancha, malim (Uzbek).

Estrus: (C) five days (Schauenberg 1979).
Gestation: (C) 63-68 days (Green 1991).

Description
and Behavior (Plate 10)
Like the serval, the jungle cat has long legs and a slenderbuild. The fur is generally sandy brown, reddish or grey,
and is unpatterned except for stripes on the legs and occasionally the throat, which are very light in the south of its
range and darker in the north (Pocock 195 1, Heptner andSludskii 1972, Harrison and Bates 1991). The winter coatis darker and denser than in summer (Heptner and Sludskii
1972). Melanistic individuals are occasionally reported(Pocock 1939a, Chakraborty et al. 1988, T. Roberts in litt.
1993). Jungle cats have black ear tufts (up to 15 mm inlength: Roberts 1977). The tail is relatively short, averaging about 40% of head-body length (TL=27 cm; n=49:
Pocock 195 1). Males are markedly larger than females(6.1 k 1.5 kg [n=20] vs. 4.2 t 1.1 kg [n=l2]: Pocock 195 1).
An old male captured in Russia’s Astrakhan reserve
weighed 13 kg (Heptner and Sludskii 1972). In captivity,
males are very protective of the cubs, even more so thanfemales, and sexual dimorphism may be linked to thisbehavior (Schauenberg 1979, H. Mendelssohn in Zitt.
1991). Family groups- male, female and cubs-havebeen seen in the wild (Schaller 1967, Mendelssohn 1989).
Allayarov (1964) described two jungle cat dens foundalong rivers in Uzbekistan: small hollows in dense reed

Litter size: (C) 2.89 (n=82); range l-6.
Interbirth interval: (C) 93- 13 1 days (Schauenberg 1979).
Age at sexual maturity: (C) 11 months (Schauenberg
1979) - 18 months (Petzsch 1968).
Longevity: (C) up to 14 years (Green 1991).
Habitat and Distribution
The jungle cat, despite its name, is not strongly associatedwith closed forest, but rather with water and dense vegetative cover, especially reed swamps, marsh, and littoraland riparian environments.
It is able to satisfy theserequirements in a variety of habitats across a wide geographic area (Fig. 16). In sandy and stony desert country(sometimes with only very sparse shrub cover: Roberts1977), it occurs along riverbeds or near oases (Heptner andSludskii 1972, Osborn and Helmy 1980, Harrison andBates 199 1, Belousova 1993, E. Matjuschkin in litt. 1993).
In southeast Asia, it is typically found in tropical deciduous forest (Lekagul and McNeely 1977, Feng et al. 1986,
Rabinowitz and Walker 1990, A. Johnsingh in litt. 1991),
although it has also been reported from evergreen forest
in central Vietnam (Trinh 1991), probably in association

83

Pat? I: Species

Accounts.

Chapter

3. TropicalAsia,

Junglecat

Figure 16. Distributionof the jungle cat (F. chaus).
1. Golardi Sulun VI; 2. Karatepe-Aslantas
V (Turkey);
3. Mesopotamian
marshes(not protected:Iraq); 4. Khoshyeylag
I; 5. Arjan IV*; 6. Khab-o-Rouchon
I (Iran); 7. Lal Suhanra V*; 8. Kirthar II (Pakistan);
9. Royal Sukla
Phanta IV (Nepal); 10. Specimen
collected in a river valley 50 km northwestof Mt. Everest (Feng et al. 1986: Tibet); 11. Gir II complex;
12. Ranthambore
II; 13. Melghat IV; 14. Nagarjunasagar
SrisailamIV; 15. Bandipur II; 16. Sanjay II complex;
17. Mouling II (India);
18. Sundarbans
E, W & S IV (Bangladesh);
19. Huai Kha Khaeng IV complex;
20. Kaeng Tana II (Thailand);21. Yok Don IV (Vietnam);
22. Flood Plains II complex (Sri Lanka); 23. Borzhom
I (Georgia);
24. Kyzyl-Agach
I (Azerbaijan);
25. Badai Tugai I (Turkmenistan);
26. Astrakhan
I (Russia).

with forest clearance. It does not occur south of theIsthmus of Kra. It is also found in shrub and grassland. It
has been recorded up to 2,400 m in the Himalayas(Guggisberg 1975), and up to 1,000 m in the Caucasus
mountains between the Black and Caspian seas
(Vereshchagin 1959). It was reported from the southeastern mountains of Algeria in the 1930s (3,000 km from the
Nile River Delta in Egypt, the only place it is known tooccur in Africa), but the skin, purchased in a market, waslater identified by Pocock (195 1) as an African wildcat(Kowalski and Rzebik-Kowalska
199 1).
Jungle cats have adapted well to irrigated cultivation,
having been observed in many different types of agricultural and forest plantations throughout their range, withsugarcane frequently mentioned in tropical Asia (Tikader
1983, Khan and Beg 1986, U. Karanth in litt. 1991, 1993).
In Israel, they are commonly found around pisciculture
ponds and irrigation ditches (Mendelssohn1989).
Vereshchagin (1959) noted that the cats’ use of the semiarid plains of Azerbaijan increased with development of a
local irrigation system and decreased with its abandonment. However, mowing the seasonally flooded riverine
tugai vegetation (trees and shrubs with dense stands oftall reeds and grasses) of this region for livestock fodder, as

well as plowing it under for agriculture, is known to be
associated with the decline of jungle cat populations insome parts of central Asia (Amudarya,
Dagestan,
Kalmykia, Karakalpakiya,
Khorezm Oasis, northern
Osetia and Syrdarya: Heptner and Sludskii 1972,
Nuratdinov and Reimov 1972, Esipov 1983, Korneev andSpasskaya 1983, Kuryatnikov 1983, Belousova 1993).
Jungle cats are often spotted amidst human settlement(and are frequently reported to take chickens). Pocock(1939a) reported that jungle cats in Kashmir occupied“nearly every old building about Srinagar,” and recently,
in southern India, a breeding pair was found occupying an
old building in an urban area, near coconut palm plantations (U. Karanth in Zitt. 1991).
Population Status
Global: Category 5b. Regional (Tropical Asia): Category
4. Regional (N Africa & SW Asia): Category 5a. IUCN:
not listed. The species is widely considered common, andis probably uncommon only in countries at the edge of its
range, such as China (Tan 1984, Wang and Wang 1986,
Gao et al. 1987). In Sri Lanka as well, Phillips (1935)
described the jungle cat as uncommon, and confined tothe dry, open country of the north.

84

Part I: Species

Density estimates from natural tugai habitat in central
Asia range from 4- 15 individuals per 10 km2 (Belousova
1993), but where this vegetation type has declined due todevelopment density does not exceed two cats per 10 km2(Nuratdinov and Reimov 1972).

Accounts.

Chapter

3. TropicalAsia,

Leopardcat

1939a, Gao et al. 1987). There is a good deal of variationin the pelage: for example, Heptner and Sludskii (1972)
describe the leopard cat in Russia, with its tiny range, as
having greater color variation than any other Soviet felid
except the lynx, which has a much wider range. The leopard cat’s pelt is dotted with dark spots which are sometimes solid, sometimes rosettes, and the tail is banded withblack rings toward the tip. Males (3.3-4 kg: n=6) are larger
than females (2.5-3 kg: n=2) (Izawa et al. 1991,
Rabinowitz1990). Male leopard cats in Russia haveweighed up to seven kg (Heptner and Sludskii 1972).
Rodents form the principal prey (China: Wang Peichao,pers. comm to A. Abdukadir;Japan: Inoue 1972;
Philippines: Alcala and Brown 1969; Russia: Stroganov
1962, Heptner and Sludskii 1972; Thailand: Rabinowitz1990). The diet also includes young ungulates, hares,
birds, reptiles, insects, eels, and fish, as well as occasional
carrion (Heptner and Sludskii 1972, Santiapillai andSuprahman 1985, Gao et al. 1987, Yu and Wozencraft inpress). Although often described as primarily nocturnal,
four radio-collared leopard cats in Thailand were frequently active during the day, and times of peak activity
varied individually (Rabinowitz 1990).
The taxonomic status of the leopard cat is controversial,and needs re-examination, with the Iriomote cat (see next
account) being the best example. Is the leopard cat a single species with pronounced geographic variation(Wozencraft 1993, Yu and Wozencraft in press), or has
isolation, particularly on islands, been sufficiently lengthy
to warrant species recognition for some populations?
Rabor (1986) has suggested that the leopard cats of Panay,
Negros, and Cebu, which are separated from the Sunda
Shelf by deep water channels, may be a different andendemic subspecies of the Philippines in comparison withthe population found on Palawan, which would be
expected to have a closer relationship to Indonesian island
populations (C. Groves, W. Oliver in litt. 1993). Yu andWozencraft (in press) recognize the leopard cats of Java,
Sumatra, Borneo, and Hainan as distinct subspecies, but
not the cats of the Philippine Islands, which have not yet
been described. Meanwhile, Heptner (197 1) has argued
that the leopard cat of northeastern Asia (Amur cat, F.b.
euptiha) should be considered a separate species, but hecompared it to leopard cats from southeast Asia and India.
When compared to Chinese leopard cat populations, his
distinctions do not hold (Gao et al. 1987).

Protection Status
CITES Appendix II. National legislation: protected overpart of its range. Hunting prohibited: Bangladesh, China,
India, Israel, Myanmar, Pakistan, Tajikistan, Thailand,
Turkey. No legal protection outside protected areas:
Bhutan, Georgia, Laos, Lebanon, Myanmar, Nepal, Sri
Lanka, Vietnam. No information: Afghanistan, Armenia,
Azerbaijan,
Cambodia, Egypt, Iran, Iraq, Jordan,
Kazakhstan, Syria, Turkmenistan, Tajikistan, Uzbekistan(IUCN Environmental Law Centre 1986, Nichols et al.
1991; A. Bukhnicashvili, U. Ohn, R. Salter, S. Umar in Zitt.
1993).
Principal Threats
Jungle cats do well in cultivated landscapes (especiallythose that lead to increased numbers of rodents) and artificial wetlands. However, reclamation and destruction ofnatural wetlands, ongoing throughout its range but particularly in the arid areas (Dugan 1993), still pose a threat tothe species, as density in natural wetlands is generally
higher (Allayarov 1964, Belousova 1993).
Action Planning
Project 80.

Leopard cat, Prionailurus
bengalensis (Kerr, 1792)
Other Names
Bengal cat (English); chat leopard du Bengale (French);
Bengalkatze (German); gato bengali, gato de Bangala(Spanish); chita biral, ban biral (Bangladesh); jin chienmao, bao mao, shih hu, shan mao (Chinese); psk jangley(Dari: Afghanistan);kucing batu, kucing congkok(Indonesia); chita billi (India, Pakistan); nam laniao(Kachin); huli bekku (Kannada: India); kla hla (Karen,
Talain); sua meo, sua pa, sua nak (Laos); wagati (Mahratti,
Ghats: India); kucing batu, rimau akar (Malaysia); kye thit,
thit kyuk, kya gyuk (Myanmar);maral, tamaral(Philippines); Amurskii kot, bengalskaya koshka (Russia);
hen wap (Shan); maew dao (Thailand).

Biology
Reproductive season: (W) Breeding is reported to takeplace once annually in the north of its range (FebruaryMarch) (Ognev 1935, Stroganov 1962, Roberts 1977,
Prater 197 1); in the tropics, year-round (Lekagul andMcNeely 1977, Santiapillai and Suprahman 1985, Gao et
al. 1987).

Description
and Behavior (Plate 10)
Leopard cats tend to yellowish-brownin the tropics andgreyish-brown in the northern parts of their range (Pocock

85

Part I: Species

Accounts.

Chapter

3. TropicalAsia,

Leopardcat

Figure 17. Distributionof theleopardcat (I? bengalensis).
1. Naltar IV (Pakistan);
2.
DachigamII; 3. KedarnathIV;
4. Bandipur II; 5. Kanha II;
6. Balphakram
II; 7. Mouling II(India); 8. Royal Sukla Phanta
IV; 9. Royal Chitwan II**
complex (Nepal); 10. Huai
Kha Khaeng IV complex;
11. Tarutao II (Thailand);12. Cut Phuong II; 13. Bach
Ma Hai Van II; 14. Nam Bai Cat
Tien II (Vietnam);
15. Lomphat
reserve (proposed:
Cambodia);
16. Taman Negara II (Peninsular Malaysia);
17. Tabin VIII;
18. Kinabalu II (Sabah, Malaysia); 19. Kutai II (Kalimantan,
Indonesia);
20. Meru Betiri II;
21. Ujung Kulong II (Java,
Indonesia);
22. Berbak IV;
23. Gunung Leuser II* complex (Sumatra,Indonesia);
24. Sundarbans
E, W and S
IV (Bangladesh);
25. St. Paul
SubterraneanRiver II (Palawan,
Philippines);
26. Panay Mts. NP(proposed:
Panay, Philippines);
27. Calauit Island IV (Philippines); 28. Tawu Mts. IV(Taiwan); 29. TsushimaIslands
protected area (Japan); 30. Mt.
Paekdu IV* (North Korea);
31. Changbai
Mts IV* (China);
32. Kedrovaya
Pad’ I (Russia).

coniferous forest, as well as shrub forest and successional
grasslands (Heptner and Sludskii 1972, Lekagul andMcNeely 1977, Santiapillai and Suprahman 1985, Fenget al. 1986, Cai et al. 1989a, Ha Dinh Due in litt. 199 1, T.
Roberts in litt. 1993). The northern boundaries of its range
are limited by snow cover; the leopard cat avoids areas
where snow is more than 10 cm deep (Formozov 1946).
It is not found in the cold steppe grasslands (Ognev 1935),
and generally does not occur in arid zones, although there
are a few records from relatively dry and treeless areas inPakistan (Roberts 1977). Leopard cats usually live in
proximity to a water source (Gao et al. 1987), and canoccupy refuge strips of riverine forest in areas otherwise
deforested (Johns 1989: 99). They are arboreal to someextent: in Thailand, one cat was radio-located resting in a

Gestation: (C) 56-70 days (Nawa 1968, Hemmer 1976).
Litter size: (C,W) l-4, usually 2-3 (Eisenberg
to eight (Heptner and Sludskii 1972).

198 1); up

Age at sexual maturity: (C) as early as eight months.
Longevity: (C) up to 15 years, but teeth lost at 8- 10 years(P. Quillen in litt. 1993).
Habitat and Distribution
The leopard cat has a wide distribution in Asia, ranging
up to 3,000 m in parts of its range, which extends into the
Himalayas along river valleys (Habibi 1977, Feng et al.
1986). It occurs in a broad spectrum of habitats, from tropical rain forest to temperate broadleaf and, marginally,86Part I: Species

tree at a height of over 20 meters (Rabinowitz 1990).
Leopard cats occur commonly in dense secondary
growth, including logged areas, and have been found in
agricultural and forest (rubber tree, oil palm) plantations(Harrison 1974, Davies and Payne 1982, Santiapillai andSuprahman 1985, M. Khan in litt. 199 I)-even breedingin hill coffee plantations in southern India (U. Karanth in
Zitt. 1993). Some have speculated that secondary forest
may be preferred to primary forest (Banks 1949,
Santiapillai and Suprahman 1985). Leopard cats can live
close to rural settlements, occasionally raiding poultry, andhave recently been reported from the outskirts of Beijing,where they were thought to have disappeared years ago(Tan Bangjie in litt. 1991).
Leopard cats are excellent swimmers (the type specimen was caught swimming in the Bay of Bengal [Pocock
19 17]), and have successfully colonized offshore islands
throughout their range. They are found on small islands
off South Korea (Japan: Tsushima islands; S. Korea: Cheju
Island), Sumatra (Santiapillai and Suprahman 1985),
Thailand (Legakul and McNeely 1977, Thailand Royal
Forest Dept. in Zitt. 1993), Vietnam (R. Cox, pers. comm.),
China (Lu and Sheng 1986), and India (Wildlife Instituteof India unpubl. data). Small islands with leopard cat populations are shaded black in Fig. 17.
Two radiotelemetry studies have produced the first dataon home range size for leopard cats, although densities
have not been estimated. On the Tsushima islands, Izawa
et al. (199 1) reported average home ranges of 0.83 km2 forfive males and one female. In dry tropical forest in
Thailand, home range sizes for three males and one female
ranged between 1.5-7.5 km2, with core areas of 0.7-2 km2(Rabinowitz 1990).

Accounts.

Chapter

3. TropicalAsia,

Leopardcat

ence of the Parties voted to downlist P.b. bengalensis toAppendix II for all range countries except Bangladesh,
India and Thailand, whose populations remain on
Appendix I. National legislation: protected over part of its
range. Hunting prohibited: Bangladesh, Hong Kong, India,
Indonesia, Japan, Malaysia (except Sabah), Myanmar,
Nepal, Pakistan, Russia, Thailand, Taiwan. Hunting andtrade regulated: South Korea, Laos, Singapore. No legal
protection outside protected areas: Bhutan, Brunei, China,
Philippines, Vietnam.
No information: Afghanistan,Cambodia, North Korea, (Nichols et al. 1991; U. Ohn, R.
Salter in Zitt. 1993; A. Amirkhanov, pers. comm.).
Principal Threats
The leopard cat appears to be more tolerant of deforestation and habitat alteration than other Asian felids, withthe exception of the jungle cat. However, it is not invulnerable, as attested to by population declines on small
islands (Izawa et al. 199 1). Captive breeding programs arebeing developed for the populations on Tsushima (Japan:
T. Doi in Zitt. 1993) and Negros islands (Philippines: E.
Alcala, pers. comm.).
In China, the center of its range, commercial exploitation has been heavy, especially in the southwest (YuJinping in Zitt. 1993). Exports from China jumped in 1984,
averaging roughly 200,000 skins annually through 1989
(WCMC, unpubl. data). The actual harvest is much higher:
a 1989 survey of major Chinese fur companies revealed
estimated stockpiles of over 800,000 pelts (Yu andWozencraft in press). While harvests of leopard cat havebeen high in the past, averaging 150,000 annually from
1955- 1981 (Lu and Sheng 1986), the annual take from1985-1988 is believed to be of the order of 400,000 (YuJinping in Zitt. 199 1). The European Community, formerly
the primary destination for leopard cat pelts exported from
China, imposed an import ban in 1988, and Japan became
the main consumer, at a lower level, importing 50,000 skins
in 1989 (Johnson and Fuller 1992). There is also a substantial domestic market (Johnson et al. 1993). Concern
over the situation has grown. In April 1993, CITES called
on Parties to refrain from importing leopard cat products
from China until it had implemented a series of recommendations to control and manage the trade. A project toinvestigate the status of the species and to advise theChinese government on the design of a sustainable management program is underway (Johnson and Fuller 1992,
Johnson et al. 1993).
Leopard cats can hybridize with domestic cats, as isshown by the popular domestic breed, the “safari cat.”Hybridization in the wild has been reported (Heptner andSludskii 1972).

Population Status
Global: Category 5b. Regional: Category 5b. IUCN: notlisted. Leopard cats are common (relative to other felids)
across much of their range. Island populations are themost vulnerable. In the Philippines, where the current
state of the forests is arguably the worst in tropical Asia
(Collins et al. 1991), the leopard cat is certainly in trouble(Cox 1988). It has perhaps been extirpated from Cebu,
which is largely deforested, and has probably been eliminated from most of its former range on other islands (W.
Oliver in Zitt. 1993). On the Tsushima islands, leopard cats
are estimated to number less than 100, down from perhaps 200-300 individuals in the 196Os-1970s (M. Izawa
in Zitt. 1991). On Taiwan, they are seldom caught in thetraps set by aboriginal hunters (Nowell 199 1).
Protection
StatusCITES Appendix II; P.b. bengalensis CITES Appendix I;
In 1985 the Chinese population of P. b. bengalensis wasdowngraded to Appendix II. In 1994 the CITES Confer-

Action Planning
Project 13.

87

Part I: Species

Accounts.

Chapter

3. TropicalAsia,lriomotecatal. 1994). The latter two studies suggest that the Iriomote
cat separated from the leopard cat less than 200,000 years
ago, which coincides with the geological isolation of the
Ryuku archipelago. Neither study made recommendations
on whether science should consider the Iriomote cat a full
species or a leopard cat subspecies, noting only that the
Iriomote cat has evolved some unique morphological characters compared with the mainland leopard cat.
The repercussions that classification of the Iriomotecat has for its conservation make it an extremely importantcase study for reconciling the results of molecular andmorphological analyses (see discussion of this issue in the
research chapter). Is it the world’s most endangered cat
species, or rather a distinctive island population of one of
the world’s most common cats? A project is put forward
in Part III to resolve this conundrum.
The Iriomote cat has a dusky brown pelt with rather
long hair, patterned with horizontal rows of darker spots
which tend to form indistinct bands. It has a relatively
elongate and low-slung build, with short legs and tail(TL=19 cm, 32% of head-body length [n=3]: Imaizumi1967). Average weight is 4.2 t 0.5 kg for males (n=15),
and 3.2 t 0.3 kg for females (n=lO) (Izawa et al. 1989).
The diet of the Iriomote cat has been studied in detail(Yasuma 198 1, 1984, 1988): 95 prey species were identified from 849 scats. Major prey species include the common rat (36%), Ryuku flying fox (16.5%), birds(brown-earedbulbul and banded crake: 7.4%), andKishinoue skink (18.6%). A variety of insects are frequently eaten (including 39 species of beetle), but they
contribute little to the diet by weight (Yamaya and Yasuma1986). Amphibians, crabs, and fish are occasionally taken.
The Iriomote cat is primarily nocturnal, partially arboreal,and swims well (Yasuma 1981).

lriomote cat, Prionailurus
bengalensis iriomotensid
lncertae sedis (Imaizumi, 1967)
Other Names
Chat d’hiomote (French); Iriomote-Katze (German); gatod’hiomote (Spanish); Yameneko (Japan).
Description
and Behavior (Plate 10)
The Iriomote cat was first described for science in the late1960s when Imaizumi (1967) labelled it “probably one of
the most primitive species among the Asiatic Felidae.” It isfound only on Iriomote island, an island of 293 km2 at thesouthernmost tip of the Ryuku chain, located about 200 km
east of Taiwan. Imaizumi (1967) considered the Iriomote
cat to be a highly differentiated form based upon the following principal characters: the ventral border of theparaoccipital process separated from the auditory bulla; anoval disc on each side of the basisphenoid and basioccipital region; P3 with postero-external comer evenly rounded,without a cingulum cusp; and auditory bullae unusuallysmall. These characters, however, are polymorphic in theleopard cat (Petzsch 1970, Corbett and Hill 1993).
The Iriomote cat is currently classified as a subspeciesof the leopard cat (P. b iriomotensis: Wozencraft 1993,
Yu and Wozencraft in press), albeit one “subjected to
extreme selective pressure with the attendant possibility ofgenetic drift” (Glass and Todd 1977). However, it was
originally described as a monotypic species (MayaiEurus
iriomotensis: Imaizumi 1967), and is also known as a
species closely related to the leopard cat (Prionailums iriomotensis: Hemmer 1978a, Leyhausen 1979, Corbett andHill 1993). Moreover, based on skull characters, close
relationships have been suggested between the Iriomote
cat and the Asiatic golden cat, the Bomean bay cat (Groves
1982), and the marbled cat (Leyhausen and Pfleiderer
1994). Leyhausen and Pfleiderer (1994) also state that
the Iriomote cat has incompletely sheathed and semiretractile claws, resembling the fishing and flat-headed
cats. They maintain that the Iriomote cat has more in common morphologically with other cats of the three genera
Prionailurus, Profilis, and P&dofelis than with the leopard cat populations of east and southeast Asia. According
to their analysis, the Iriomote cat is most properly classified as Prionailurus iriomotensis, although further investigation may well lead to the resurrection of the original
genus Mayailurus.
However, while Leyhausen and Pfleiderer (1994) argue
for a relatively distant relationship between the leopard cat
and the Iriomote cat based on morphological characters,
molecular analyses have led the investigators to conclude
that the leopard cat is the Iriomote cat’s closest relative(Wurster-Hill
et al. 1987, Masuda et al. 1994, Suzuki et

Biology
Reproductive season: (W) Mating in February-March and
September-October,based on vocalizations.
Birthsobserved only late April-May.
Gestation: (W) approx. 60-70 days.
Litter size: (W) l-4 (Yasuma 1984, 1988).
Longevity: (C) One male kept by the Okinawa Kodomonokuni Zoo died at an age of over 10 years (T. Doi inlitt. 1993).
Habitat and Distribution
Found only on Iriomote Island, at the southernmost tip of
the Ryuku Island chain, which is part of the archipelago
stretching from Kyushu to Taiwan known as Nansei Shoto(Fig. 18). Iriomote Island consists predominantly of low
mountains (300-400 m) covered with sub-tropical evergreen broadleaved primary forest, including extensive belts
of mangrove along waterways.
The Nansei Archipelago
88Pat? I: Species

Accounts.

Chapter

3. TropicalAsia,lriomotecatshowed higher residentiality, maintaining stable homeranges and using one feeding site for several years, whilemales tended to shift their area of activity after a period ofseveral months. In general, male home ranges overlapped
those of other males and females, while female home ranges
seldom overlapped (Izawa et al. 1989). Density is approximately 0.34 animals per km2 (M. Izawa in Zitt. 1993).

CHINA

habitatofcat (I 200m)

tIJ

lriomote
National

II
Park

q

Montanearea whereoccur at low density

I3

Main

lriomotecats

Highway

Taiwan//FigurePreferred
lriomote

Protection
StatusCITES Appendix II. National legislation: fully protected
since 1967, but new endangered species legislation setsstronger requirements for habitat protection, and thenational government is now investigating ways to expand
protected area coverage (M. Izawa in Zitt. 1993). The park
does not protect the most important habitat for the Iriomotecat, lowland coastal forest (Izawa et al. 1991: see Fig. 18).
However, the Japanese government is in the process ofdeveloping a comprehensive action plan for both researchand conservation (T. Doi in Zitt. 1993). The best prospect
for further reserve establishment is along the western
coastline, but the area has not yet been studied to determine its suitability for the Iriomote cat (M. Izawa in Zitt.
1994), and creating more reserves on Iriomote will bepolitically difficult.

p L lshigaki
Islandtlriomote
Island

18. Distributionof the lriomote

Principal Threats
The Iriomote cat is endangered primarily because it isrestricted to a single population, albeit located on the least
developed island in Japan. That situation is changing
rapidly, however, as island residents press for accelerated
economic growth. Iriomote is promoted as a tourist location, with the Iriomote cat a major source of appeal. The
industry is still nascent, but plans are being laid for major
resort development, along with a dam to provide the eightprojected hotels with a stable supply of water (Anon.
1992a). A major airport is being constructed on nearby
Ishigaki island (20 km away) to provide a direct link toTokyo.
While poorly planned tourist infrastructure may damage the cat’s habitat, by far the major threats at present
are agricultural and cattle-raising projects, which are heavily subsidized by the government (Barber et al. 1984, Anon.
1992a) and involve forest clearance. Conservationists’opposition to the Ministry of Agriculture’s projects andtheir calls for legal protection of lowland habitat have further increased the local residents’ impression that the
Iriomote cat is an obstruction to economic development.
Other important threats include road kills, competition
from a growing population of feral cats, and the risk of disease transmission from these and other imported mammals(Izawa et al. 1991, M. Izawa in litt. 1993).

cat (P. b. iriomotensis).

has a high degree of endemism, and has been termed the“Galapagos of the Orient.” People came to the island only
after the second World War, following spraying of DDT by
the Americans to eradicate malaria (P. Leyhausen in Zitt.
1977), and by 1991 numbered around 1,500 (Izawa et. al.
1991). Settlement is concentrated mainly along the coastfrom the northwest to the southeast, with the center of theisland being relatively undisturbed (Barber et al. 1984,
Anon. 1992a). However, the Iriomote cat typically inhabits the low altitude coastal areas of the island alongside
the human population, rather than the protected montane
area at the island’s center (Izawa et al. 1989).
Population Status
Not ranked here for vulnerability, but with its tiny range
and small population size, the Iriomote cat as a full species
would be in Category 1, and qualify as the world’s rarest
and most vulnerable cat. IUCN: Endangered. The population is estimated to number less than 100 individuals, but
is thought to have remained stable since monitoring began
in 1982 (Izawa et al. 1989).
Radiotelemetry studies carried out from 1982- 1988
found that males had larger home ranges than females, at
2.96 2 1.8 km2 and 1.75 2 0.8 km2 respectively. Females

Action Planning
Projects 69 and 70.

89

Part I: Species

Accounts.

Chapter

4. Eurasia

Part I
Species Accounts

Chapter 4
Eurasia

Box 1
Vulnerability

Index to Species of the Region (in orderof vulnerability)

Asia sub-region
Species

Habitat Association
St [Mar] (Tot) Score

Geog.
Score
Range
(I@ kma)

Body
Size
Score

Total
Score

Ranking

Snow leopard, U. uncia*
Chinese mtn. cat, F. bieti”
Manul, 0. rnanul”
Asiatic wildcat,
F.s. ornata group
Eurasian lynx, L. /ynx*

I: 1 [6]N: 2 [3]1:4[2]

-1
+I
+I

-2
-2
+I

1 (A)
1
2

+I
+2

2
3

(7)
(5)
(6)

0
4
0

s:
R:
M:

2.39
0.29
5.08

-1
-2
0

L
s
s

l:3[4](7)
B: 6 [5] (11)0+I

M: 7.00
W: 12.61

0
+I

S+l
MO

Key:
* Most or all of this species’ range lies within the region

Habitat Association
St = number of strong + significant habitats
N = Narrow (-1); B = Broad (0)[Mar] = number of marginal habitats(Tot) = total number of habitats

Geographic Range(in millions of km*)
R = Restricted (-2); S = Small (-1); M = Medium(0); W = Wide(+1)

Body Size
L = Large (-1); M = Medium(0); S = Small (+I)

(A) = Actively threatened

Regional Criteria:
Habitat association: Narrow = 5 habitat types; Intermediate = 6-7 habitat types; Broad = 11 habitat types
Geographic range: Restricted = -<I million km*; Small = 2-3 million km*; Medium = 5-7 million km*;
Wide = 12-I 3 million km*
Body size: Large = 35-135 kg; Medium = 7-20 kg; Small = 26.5 kg

90

Pat? I: Species

Accounts.

Chapter

4. Eurasia,

SnowleopardEurope sub-region
Score

Body
Size
Score

Total
Score

Ranking

0

-2

0

-1

1
2

0

3

Species

Habitat Association
St [Mar] (Tot) Score

Geog.
Range
(106 km*)lberian lynx, L. pardi/~s*
European lynx, L. /ynx
European wildcat,
Fs. silvestr& group

I:3
I: 5

[3](6)[I] (6)

0
0

R:
S:

0.08
0.95

-2

M

-1

M

I:4

[2](6)

0

S:

1.67

-1

s

+I

* All of this species’ range Iies within the region
Regional Criteria:
Habitat association: Intermediate = 6 habitat types
Geographic range: Restricted = co.5 million km2; Small = 0.52 million km2
Body size: Large = 35-135 kg; Medium = 7-20 kg; Small = 16.5 kg
See the Introduction to the Species Accounts for explanation of the vulnerability ranking system (pp. Z-6).

little from the winter in density and length (Heptner andSludskii 1972). The long tail is thought to aid balance,and
snow leopards will wrap their tails around themselves
when lying or sitting for addedwarmth. The snow leopard’s vocal fold is lessdevelopedthan in the pantherines,
lacking a thick pad of fibro-elastic tissue,so that it cannot
make the low and intense “roars” of which the other big
cats are capable(Hemmer 1972,Peters1980,Hast 1989).
Males are larger than females, with average weights
between 45-55 kg as opposed to 35-40 kg for females(Jackson1992).
A. Kitchener (in Zitt. 1993)hassuggestedthat the snow
leopard, for which two subspecieshave been described(Stroganov 1962) but are not generally recognized (e.g.,
Hemmer 1972,Wildt et al. 1992a),is a prime candidateforsubspeciationbecauseof the insular and patchy nature of
its high mountain habitat. Similarly, Fox (1994) draws
attention to the gapbetweenthe main southernsnowleopard population and the northern population in Russiaand
Mongolia, and suggeststhat the two populationsmay differ genetically. On the other hand,instancesof snowleopardsmigrating up to 600 km have beenreported from theformer U.S.S.R. (Heptner and Sludskii 1972, Koshkarev
1990: Fig. 1).
Snow leopardsare opportunistic predatorscapableof
killing prey up to three times their own weight (Schaller
1977, Jackson and Ahlborn 1988, Fox 1989), with the
exception of fully grown yak or wild ass. They will also
take smallprey: in China’s Qinghai province, Schaller etAsia Sub-region
Snow leopard, Uncia uncia(Schreber, 1778)
Other Names
Ounce (English); pantheredesneiges,leoparddesneiges,
once (French); Schneeleopard,Irbis (German); leopard0
nival, panterade las nieves (Spanish);xue bao (Chinese);palang-i-berfy (Dari: Afghanistan); bharal he, barfani
chita (Hindi, Urdu: India, Pakistan);shan(Ladakhi: India);hiun chituwa (Nepal); lkar (Pakistan); irbis, irvis (Russia,
Central Asian republics, Mongolia); snezhnai bars
(Russian);sarken(Tibetan); then (Bhutan).
Description and Behavior (Plate 11)
The snow leopard exhibits superb camouflage for its
mountain environment of bare rocks and snow, being
whitish-grey (tinged with yellow) in color, and patternedwith dark grey rosettesand spots. Further adaptationsfor
high altitude life include an enlargednasalcavity, shortenedlimbs (adult shoulder height is about 60 cm), welldeveloped chest muscles(for climbing), long hair with
dense,woolly underfur (belly fur grows aslong as12 cm),and a tail up to one meter long, 7590% of head-bodylength (Hemmer 1972, Fox 1989, Jackson 1992). Snowleopardsmolt twice a year, but the summercoat differs

91

Pat? I: Species

Accounts.

Chapter

4. Eurasia,

Snowleopardstock despite the availability of blue sheep in relatively
high numbers (Oli 1991).

al. (1988a) found that 45% of their summer diet consistedof marmots. In general, their most commonly taken prey
consists of wild sheep and goats (including blue sheep,
Asian ibex, markhor, and argali), but also includes pikas,
hares, and gamebirds (chukor partridge and snowcocks)(Hemmer 1972, Heptner and Sludskii 1972, Schaller
1977, Jackson 1979, Mallon 1984a, Schaller et al. 1987,
1988a, Fox 1989). Jackson and Ahlborn (1984) estimateda snow leopard’s annual prey requirements to be of theorder of 20-30 adult blue sheep. In a study of radio-collared animals, Jackson and Ahlborn (1988) found that
adult snow leopards killed a large prey animal every lo- 15days, and remained on the kill for an average of 3-4 days,
and sometimes up to a week. Snow leopards tend toremain within a relatively small area for 7- 10 days, thenshift activity to a relatively distant part of their homerange. Daily distances moved were up to seven km, but
averaged one km for males and 1.3 km for females(Jackson and Ahlborn 1988).
Predation on livestock can be significant (Schaller
1977, Mallon 1984a, Fox and Chundawat 1988, 1991a,
Schaller et al. 1988a,b, Chundawat and Rawat 1994, Oli
1994, Jackson et al. 1994), with stock losses on the Tibetan
Plateau averaging about 2% per village, but up to 9.5% insome “hotspots” (Jackson et al. 1994). Oli et al. (1994)
analyzed 2 13 scats of snow leopards living around villages
within Nepal’s Annapurna Conservation Area, and foundlivestock remains in 17.8%. The proportion increased to
39% in winter, probably in relation to marmot hibernation,
deep snow, and a tendency for yak to be less widely dispersed at this time. Snow leopards in this area took live-

Table 1
Distribution

and Population

Estrus: (C) 2-12 days (Rieger 1984).
Estrus cycle: (C) 15-39 days (Freeman 1975).
Gestation; (C) 98-104 days (Jones 1977, Freeman 1975).
Litter size: (C,W) l-5, usually 2-3, exceptionally up toseven (Heptner and Sludskii 1972, Blomqvist and Sten
1982, Wharton and Freeman 1988).
Age at dispersal: (W) 18-22 months; sibling groups may
remain together briefly upon independence (Jackson andAhlbom 1989).
Age at sexual maturity: (C) 2-3 years.
Age at last reproduction: (C) 15 years.
Longevity: (C) up to 21 years (Blomqvist
Wharton and Freeman 1988).

and Sten 1982,

Habitat and Distribution
As shown in Fig. 1, the snow leopard has an extremely
patchy and fragmented distribution, consisting of a mix oflong narrow mountain systems and islands of montane

Estimates for Snow Leopard (FOX 1994)

Area of

Estimated

Habitat (km*)

Population

Afghanistan
BhutanChina
India
Kazakstan
Kyrgyzstan
Mongolia
Nepal
Pakistan
RussiaTajikistan
Uzbekistan

50,00015,0001,100,000 ’
75,00050,000105,000
90,00030,00080,000130,000
100,000
to,000100 - 200
100 - 200
2,000 - 2,500
200 - 600
I&O - 200
800 - 1,400
500 - 1,000
350 - 500
100 - 200
50 - 150
120 - 300
IO - 50

Total

1,835,OOO

4,510 - 7,350

Country

Biology
Reproductive season: (W) early January to mid-March, atime when vocalizations can most commonly be heard(Jackson and Ahlborn 1988). (C) same; most births occur
in May-June (Freeman 1975, 1977, Blomqvist and Sten1982).

literature

SourceMap-based estimates (low density)
Map-based estimates (moderate density)
Schaller 1990, Jackson 1992
Chundawat et al. i988, Fox et al. 199laAnnenkov 1990, Zhirjakov 1990
Zhirjakuv 1990, E. Koshkarev, pers. comm.
Green 1988, Schaljer et al. I994
R. Jackson, pers. comm.
Schaller 1976, 1977
Smirnav et al. 1990, E. Koshkarev, pers. comm.
Sokov 1990, Buzurukuv, pers. comm.
Braden 1982, E. Koshkarev, pers. comm.

92

Pat? I: Species

qn

Protected
area of sufficient
a minimum
viable population

size to conserveof 50+ breeding

Accounts.

adultsL

Figure 1. Distributionof the snow leopard(U. uncia) after Fox (1994).
1. Sayano-Shushenskiy
I* (Russia); 2. Pamir-i-Buzurg
IV (Afghanistan);
4. Khunjerab
II (Pakistan);
3. Taxkorgan
IV (Xinjiang, China) + Khunjerab
II (Pakistan);
4. Hemis II (India); 5. Shey-PhoksundoII
complex;6. Annapurna
Conservation
Area (Nepal); 7. Qomolongma
IV (Tibet, China) + Langtang
II+ Sagamartha
II** (Nepal) complex;
8. Jigme Dorji IV (Bhutan);
9. Adult snow leopard captured in thewinter of 1957-1958
on the shore of Lake Balkhash,more than 600 km from the nearest mountain
ranges (Heptnerand Sludskii 1972).

93

Chapter

4. Eurasia,

SnowleopardPat? I: Species

Accounts.

Chapter

4. Eurasia,

SnowleopardTable 2
Protected Areas for Snow Leopards by Range State and Size
(International

Snow Leopard

Trust, May 1993)

Country< 500 km*500-I ,000 km*

Afghanistan
BhutanChina
India
Kazakhstan
Kyrgyzstan
Mongolia
Nepal
Pakistan
RussiaTajikistan
Uzbekistan

0
3
24
0
4
0
2
18
1
2
1

Total

56

Jackson (1992) stresses
such as in key unprotected--- Number of Protectedl,OOO-6,000 km*

Areas - - -__ ___ --.--___
5,000-l 0,000 km*

1
0
3

0
0
5
7
0
0
I?
6
2
0
0
0

0
0
0
0
0
0
0
0
0
0

19

21

1

0

the importanceof establishing
more coherent
habitat along the Mongolia-Kazakhstan-China-Russia- __I___- >I 0,000 km*

0trans-frontierprotected
border areas.

area systems,

suchasmajor ridgelines,bluff edges,gullies, andthe baseor crest of broken cliffs (JacksonandAhlbom 1988).

habitat scatteredthroughout a vast region surroundingthe
central Asian desertsand plateaus. Although the snow
leopard’srangeextendsover some2.3 million km2 of central Asia, occupiedhabitat is estimatedat only 1.6 million
km”, most of which is in Tibet and other parts of China(Fox 1994). Through most of their range, snow leopards
are associatedwith steeprocky slopeswith arid and semiarid shrubland,grassland,or steppevegetation (Fox 1989,
Jackson 1992). In the mountainsof Russiaand parts of
the Tian Shan they visit in open coniferous forest, but
generally avoid denseforest (Heptner and Sludskij 1972,
E. Koshkarev, pers. comm.).
Snow leopards are generally found at elevations
between 3,000-4,500 m, although they occasionally go
above 5,500 m in the Himalaya, and at the northern limitsof their rangecanbe found between600- 1,500m (Heptnerand Sludskii 1972,Fox 1989,Schaller et al. 1994). Steepterrain brokenby cliffs, ridges,gullies,and rocky outcrops
is preferred (Koshkarev 1984,Mallon 1984a,Jacksonand
Ahlborn 1984,1988,Chundawat1990b,Fox et al. 1991a),although in Mongolia and on the Tibetan Plateauthey canbe found in relatively flat country (Mallon 1984b,Schaller
et al. 1994),especiallyif ridgesoffer suitabletravel routes,
and shrub and rock outcrops provide sufficient cover(Schaller et al. 1988a). In general,snow leopardstend to
move, bed, and mark along linear topographic features,

Population

StatusGlobal: Category 2(A). Regional (Eurasia): Categoryl(A). IUCN: Endangered. Estimates of the total snow
leopard population vary from 4,500 to 7,500 individuals(Jackson 1992, Fox 1994). Earlier lower estimates,e.g.,
1,500by Green (1988) and4,000 by Fox (1989), reflecteda lack of information from large areasof snow leopard
range. China is home to the largest number of animals(mainly in the Tibetan region), and Kyrgyzstan andMongolia hold the next largestpopulation. Country estimatesare shownin Table 1.
Status surveys for the snow leopard have been relatively extensive, compared to other species, and theInternationalSnow LeopardTrust (ISLT), basedin Seattle,
Washington (U.S.), maintainsa databaseof field reports
and local population estimates.The Trust is developing a
protocol for standardizing survey techniques (ISLT in
prep.), and hassignedagreementsto hold training workshopsin key range states(R. Jackson in Zitt. 1993). The
Trust hasorganizedthree seminarsin central Asian range
stateswhich have successivelyrefined understandingof
the snow leopard’s status, biology, and distribution(Freeman1988,Blomqvist 1990,Fox and Du 1994).

94

Part I: Species

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Chapter

4. Eurasia,

Snowleopardwork of a national conservation strategy modelled after
Project Tiger (Govt. of India 1988); however, most ofthese reserves will be less than 500 km2 in size, reflecting
the scarcity of unpopulated land in the country.
Green (1994) reviewed the status of central Asian protected areas known to contain snow leopards. In general,
the integrity of these protected areas is poor: 65% areinhabited by people, and 86% provide grazing lands forlivestock. Only 49% have management plans, some ofwhich are pending approval while others require updating.
Green (1994) emphasizes the need to focus attention on
managing snow leopard populations in unprotected lands,
noting Jackson and Ahlbom’s (1990) estimate that 65% of
Nepal’s snow leopards live outside the country’s protectedareas (see Part II Chapter 1 for more discussion of the status of protected area coverage for snow leopard range).

Estimates of snow leopard density range from 0.8 animals per 100 km2 (Koshkarev 1989, Annenkov 1990) to
lo/100 km2 (Jackson and Ahlborn 1989). Other studies
have provided density estimates for local populations
within the following countries (expressed in terms of individuals per 100 km”): Nepal-57(Oli 1991); China-O.54 (Schaller 1988a,b); India-0.5-6.6(Fox et al. 1991a);
Mongolia-4.4(Bold and Dorzhzunduy1976) to 5
(Schaller et al. 1994); Kazakhstan-0.8-4.7(Koshkarev1989); Russia-0.75-1.5(Sopin 1977). Many of theseestimates are derived from indirect sign indices (scrapes,
scats, scent sprays, and claw rakings) along trails.
Only a few radiotelemetry studies documenting homerange have been carried out (Chundawat 1990b, Schaller
et al. 1994, M. Oli, pers. comm.). Home range size of fivesnow leopards in prime habitat in Nepal’s Langu Gorge
ranged from 12-39 km2, with substantial overlap between
individuals and sexes (Jackson and Ahlborn 1989). Small
core areas (consisting of 14-23% of total home range)
were more heavily used and marked. Core areas were
not exclusive, and were used by different animals at different times.

Principal Threats
Large ungulates have been hunted out of many areas of thehigh central Asian mountains (Schaller 1977, Cai et al.
1989b, Fox et al. 1991b, Jackson 1992), and large-scalepika and marmot poisoning programs have also been conducted on the Tibetan Plateau (Smith et al. 1990, Miller
and Jackson 1994; see discussion in the next species
account). Livestock depredation tends to be greater inareas where wild sheep and goat populations have been
depleted (Miller and Jackson 1994, Schaller et al. 1994),
although prey availability is not the only factor influencing
depredation (Oli 1994, Jackson et al. in prep.). See PartII Chapter 2 for detailed discussion.
There is demand for snow leopard bones for use as substitutes for tiger bone from the Chinese medicine trade(Liao and Tan 1988). Traders will pay up to U.S. $190
for a snow leopard skeleton in Tibet (Jackson et al. 1994).
In northern Nepal, people have been seen to trade snow
leopard bones for sheep along the border with Tibet(Jackson 1992). Garments of snow leopard fur were once
highly prized in the fashion world, with high quality coatsvalued at up to U.S. $50,000 (H. Freeman, pers. comm.).
Heptner and Sludskii (1972) and Fox (1989) review central Asian and Russian exports of snow leopard skins during the 20th century; world trade was of the order of 1,000
pelts per year in the 1920s. Although no longer in intemational trade (see Table 1 in Part II Chapter 4) fur coats
have been seen for sale in shops in Kathmandu (Barnes
1989), and “novelty” furs have been seen for sale throughout China, including Taiwan (Anon. 1987e, Low 199 1,
Jackson 1992, Fox 1994), as well as Mongolia (D. Mallonin Zitt. 1994).

Protection Status
CITES Appendix I. National legislation: fully protected
over most of its range. Hunting prohibited: Bhutan (onlyin protected areas, which cover most of snow leopard
range in this country), China, India, Kazakhstan, KyrgyzStan, Nepal, Pakistan, Russia, Tajikistan, Uzbekistan.
Hunting regulated: Mongolia (although trophy hunting is
no longer permitted). No information: Afghanistan (Fox
1989, Nichols et al. 199 1, H. Freeman, pers. corm-n.).
Occurrencein Protected Areas
Protected areas with estimated populations of 50+ breeding adults are marked with a square in Fig. 1 (J. Fox, R.
Jackson in Zitt. 1993).
The International Snow Leopard Trust maintains a
database of protected areas where snow leopards are a)definitely, b) likely, and c) possibly present. The total
number of protected areas currently stands at 10 1, andcould rise to 115- 120 when updated for China andMongolia. More than half of the reserves are less than 500
km2 in size, and are likely to harbor only a few breedingpairs (R. Jackson in Zitt. 1993). The 47 protected areas
where the presence of snow leopards is confirmed add up
to 224,284 km2 (Green 1994), about 12% of the total rangeof the snow leopard (Fox 1994).
However, according to Green (1994), the protected area
network is unlikely to grow much more: while 76 more
protected areas have been proposed in the region and await
official recognition, they will add less than 25,000 km2 to
the total. Nearly all are in India and will form the frame-

Action Planning
Projects 7 l-75.

95

Part I: Species

Accounts.

Chapter

4. Eurasia,

ChinesemountaincatChinese mountain cat, Felis
bieti Milne-Edwards, 1892

nantly nocturnal, active from dusk to dawn in captivity(B. Tan in Zitt. 199 l), and hunting primarily in the earlymorning and evening in the wild (Liao 1988). They rest
and tend their young in burrows, typically situated onsouth-facing slopes. Males and females live separately,
and the burrows inhabited by females tend to be deeper
and more secure, with only one entrance (Liao 1988). Scat
analysis indicates that rodents are the major prey (90%),
primarily mole-rats, white-tailed pine vole, and pikas.
Birds, including pheasants, are also caught. Liao (1988)
observed mountain cats hunting mole rats by listening fortheir movements through their subterranean tunnels (3-5cm below the surface), and digging them out.

Other Names
Chinese desert cat (English); chat de Biet (French);
Graukatze (German); gato de Biet, gato de1 desert0 de
China (Spanish); mo mao, huang mo mao, cao shihli(Chinese); she1 misigi (Kazakh); qel mushiiki (Uygur).
Description
and Behavior (Plate 11)
The Chinese mountain cat, endemic to China, is one of
the least-known cats. It has a stocky build, with relatively
short legs. Its coat is pale grey-fawn in winter, somewhat
darker brown in the summer, and marked with indistinct
horizontal stripes on the sides and legs. Its ears have slightdark brown tufts. The tail is fairly short (35 cm: Jacobi
1923), about 40% of head-body length; it is banded with 56 dark grey bands, and has a black tip. The auditory bullae
are moderately large, measuring about 25% of total skull
length (Pocock 1951). A wild male and female brought
to the Beijing Zoo weighed nine and 6.5 kg, respectively(Tan 1984).
What little is known of this species in the wild is mainly
due to the efforts of collectors from the Xining Zoo, who
obtained 34 specimens between 1973-1985 (Liao 1988,
B. Tan in Zitt. 199 1). Chinese mountain cats are predomi-

Known

Figurespecies

Biology
Reproductive season: (C & W) January-Marchseason, litters often born in May.
Litter size: 2-4.
Age at independence: 7-8 months (Liao 1988).

Habitat and Distribution
The Chinese mountain cat is known only from the northeastern edge of the Tibetan Plateau (Fig. 2). It has beencollected most frequently from Qinghai province, but also
from the mountains of southern Gansu and northern
Sichuan. Reports of it occurring further north and east, inrange

2. Known

distribution

matingof the Chinese

96

mountaincat (F. bietl).

Part I: Species

Accounts.

Chapter

4. Eurasia,

Manuldomestic livestock for graze. Zinc phosphide was one of
the main chemicals used (G. Schaller in Zitt. 1992), fromthe onset of control efforts up until 1978, when its use wasdiscontinued because it was discovered that it also killed
carnivores that preyed on pikas. Control programs using
poisonous chemicals continue throughout much of theChinese mountain cat’s range (Smith et al. 1990), and have
eradicated pikas from large areas (A. Smith, pers. comm.
1994). However, research has indicated that pikas reach
their greatest densities and cause greatest damage whenrangeland has already been significantly degraded bydomestic stock (Shi 1983, Zhong et al. 1985), suggestingthat the authorities could most effectively control pika
populations by focusing their efforts on measures to prevent overgrazing. Healthy predator populations should
serve to limit pika numbers, as pikas are an important foodsource for a variety of carnivores and birds of prey (Smithet al. 1990).
No other threats are known. G. Schaller (in Zitt. 1992)
noted that pelts of this species can be commonly found in
markets in Xining, and Low (199 1) saw two mounted
specimens for sale in southern China. It would seem
unlikely, however, that hunting efforts specifically target
the mountain cat.

flatter, more desert-like terrain (F.b. chutuchta and F.b.
veUerosa: Pocock 195 1), probably refer respectively tomisidentified specimens of Asiatic wildcat and domesticcat (Haltenorth 1953, Groves 1980). It may occur along
the northern edge of the Tibetan Plateau, in the desert
mountains of Xinjiang (Pamir and Kunlun Mountains:
Wang and Wang 1986, X. Gao in litt. 1993), but suchreports have yet to be confirmed (Achuff and Petocz 1988,
A. Abdukadir in Zitt. 1993). The southernmost records
near Chengdu (Fig. 2) are from the same sort of area where
the giant panda is found, an entirely different habitat typeconsisting of montane bamboo forest. Allen (1938) notesthat these specimens, obtained in the late 1800s in the fur
markets of Tatsienlu and Sungpan (Sichuan province),
were probably not locally obtained, and speculates thatthey came from “the borderlands of the extreme westernedge of China or even from Tibet.”
According to Liao (1988), the Chinese mountain cat isfound throughout the Datong and Daban mountains
around Xining (where eight skins were collected by
Buchner in 1893: Groves 1980), at elevations ranging from2,800-4,100 m. It chiefly inhabits alpine meadows andscrub. It has also been found in hilly loess steppe andconiferous forest edge. Despite its traditional name
(Chinese desert cat), it appears not to be a desert cat at all(Groves 1980), although it may occur there marginally(Liao 1988; A. Abdukadir, X. Gao in litt. 1993). Chinese
specialists meeting in Beijing in 1992 concurred with
Groves’ (1980) suggestion that it be described as the“mountain cat” (Jackson 1992b).

Action Planning
Projects 76 and 80.

Manul, Otocolobus(Pallas, 1776)

Population Status
Global: Category 2. Regional (Asia): Category 1. IUCN:
Insufficiently Known. There is no information on status orabundance, and no records of occurrence in protectedareas. The Chinese mountain cat appears to have a very
limited distribution, but may have a much wider range
further west on the edge of the Tibetan plateau. It is interesting that Liao (1988) collected most of his animals frommountainous areas very close to Xining and Lanzhou, thecapitals of Qinghai and Gansu provinces.

manul

Other Names
Pallas’s cat (English); chat manul (French); Manul(German); gato manul, gato de Pallas (Spanish); yalam(Bashkir); malem (Bukharian); tu sun, wulun, manao,
yang shihli (Chinese); psk kuhey (Dari: Afghanistan);malin (Kazakhstan, Mongolia); kazail malin (Kazakh,
Ustyurt region); madail (Kyrgyzstan);ribilik (Ladakhi:
India); manul (Russia); sabanchi (Smirech’e and Kazakh);mana (Soyot); molun (Uygur); malin, dala mushugi(Uzbek).

Protection Status
CITES Appendix II. National legislation: fully protected
in China. The species is currently classified as a Category
II species under Chinese law, and the 1992 meeting of the
Cat Specialist Group in Beijing recommended upgrading
to Category I, which requires permission of national, rather
than provincial, authorities to hunt or trade.

Description
and Behavior (Plate 12)
Peter Pallas, who first described the manul, erroneously
suggested that it was the ancestor of the long-haired
Persian breeds of domestic cat, because of its long fur,stocky build and flattened face. The hair on its underparts
and tail is nearly twice as long as on the top and sides (Gaoet al. 1987). Like the snow leopard, this presumably helps
keep the animal warm when it hunts on snow, cold rock, orfrozen ground (A. Abdukadir in Zitt. 1993). The back-Principal Threats
Large-scale poisoning campaigns have been conductedsince 1958 in China in an attempt to control “pest” populations of pikas, which are viewed as competitors of97

Par? I: Species

Accounts.

Chapter

4. Eurasia,

Manulchukor partridge (Roberts 1977). Manuls are generally
crepuscular, being most frequently encountered at dusk orin early morning, but are occasionally seen at mid-day(Heptner and Sludskii 1972). They den in small caves
and rock crevices, and may take refuge in the burrows ofother animals such as marmots, foxes, and badgers(Bannikov 1954, Y. Ma, pers. comm. 1992). Heptner andSludskii (1972) reported that tame manuls hunting forrodents caught not only animals running on the surface,but also successfully ambushed them by hiding near exitsof burrows, using their paws to fish out the inhabitants
when the holes were shallow enough.

ground color of its fur varies from grey in the north of its
range to fox-red in some parts of the south (Ognev 1935,
Pocock 195 1, Roberts 1977), although greyish animals
are also found in the south (Heptner and Sludskii 1972).
The hairs have white tips, producing a silvery, frosted
appearance in all but the reddest specimens. The body iscompact, with short legs marked with indistinct blackbands, and a thick, short, black-tipped tail (about 45% ofhead-body length). Weight ranges from 2-4.5 kg (Pocock
1939a, Heptner and Sludskii 1972, Gao et al. 1987). The
forehead is patterned with small black spots. Its ears are
small and rounded and set low on the sides of the head.
The auditory bullae are enlarged, similar to those of thesand cat (Pocock 195 1, Heptner and Sludskii 1972). The
barking call of the manul is similar also to that of the sandcat (Heptner and Sludskii 1972) and, likewise, the low profile of its head is an adaptation to hunting in open country
where there is little cover (Pocock 1907b).
In the Lake Baikal region, analysis of 502 scats foundpikas to form the major part of the manul’s prey (89%),
with small rodents also frequently taken (44%). Other
prey included susliks (3%), birds (2%), and insectivores(1%) (Fetisov 1937). Bannikov (1954) reported that one
cat’s stomach from Mongolia contained the remains of 16
voles; another contained two pikas, one vole, and a hamster. Pikas and small rodents were also reported to be themajor prey in Ladakh (Stockley 1936) and China (Feng etal. 1986, Gao et al. 1987, Anon. 1987a, Cai et al. 1989a).
One cat in Baluchistan, Pakistan, was found feeding on

Biology
Reproductive season: (C & W) Most litters born AprilMay (Fetisov 1937, Bannikov 1954, B. Tan in litt. 1991).
Estrus: (C) 26-42 hours (n=l: Schauenberg 1978) - 5 days
(n=2: Mellen 1989).
Estrus cycle: (C) 46 days (n=l: Mellen 1989).
Gestation: (C) 66-67 days (n=2: Mellen 1989); 74-75 days
(n=l: Schauenberg 1978).
Litter size: (C,W) 3.57 t 0.53 (n=7: Mellen 1989); range
up to six or eight (Heptner and Sludskii 1972).
Age at sexual maturity:1989).

(C) females - one year (Mellen

Longevity: (C) up to 11.5 years (Jones 1977).

Figure 3. Distributionofthe manul(0.
mad).
1. Khoshyeylag
I; 2. Moteh
V (Iran); 3. Syunt-Khasardag+ Kopetdag
I complex;
4. Badkhyz
I (Turkmenistan);
5. Ziarat Juniper IV (Pakistan); 6. TaxkorganIV;
7. Arjin Mts. IV; 8. Boghdad
Mts. IX*; 9. Wolong IV*;
10. Qomolongma
IV (China);
11. Specimen
collected fromthe Sarai-Bulag
Mountains,near Yerevan,
Armenia(Ognev
1935).

Hlstoncal
and

range
Sokolov

(mid
1984,

- 1900’s:
A

Abdukadir

Bannikovin Mt.

1993)

98

Pat? I: Species

Accounts.

Chapter

4. Eurasia,

Asiatic

wildcating Inner Mongolia and Manchuria) in the early 1950s wasof the order of 10,000 (Tan 1984). Annual take in
Mongolia in the early 1900s was reportedly as high as
50,000 skins (Heptner and Sludskii 1972). Between 195868, harvests averaged 6,500 animals (Mallon 1985). In themid- 1970s annual harvest in Afghanistan was estimated tobe 7,000 (Rodenburg 1977). Harvests in the former Soviet
Union declined in the 1970s suggesting a decrease inabundance (Bannikov and Sokolov 1984). Harvests also
declined in China in the 197Os-1980s prior to extension oflegal protection to the species (Tan 1984). Mongolia was
the primary exporter of manul pelts in the 1980s with 9,185
exported in 1987, but hunting was prohibited in 1988, andexports have essentially ceased (WCMC unpubl. data).
Poisoning to control pika populations has taken place
on a large scale in parts of the Russian Federation (southwest Transbaikalia, Tuvinskaya, Altai Mountains), where
they are considered to be vectors for plague, and parts ofChina (Qinghai, Gansu, and Inner Mongolia), where they
are considered to compete with domestic stock for graze(Smith et al. 1990).

Habitat and Distribution
The manul is adapted to cold arid environments and has awide distribution through central Asia (Fig. 3), but is relatively specialized in its habitat requirements. It is foundin stony alpine desert and grassland habitats, but is generally absent from lowland sandy desert basins (Bannikov
1954; E. Matjuschkin in litt. 1993), although it may penetrate these areas along river courses (Ognev 1935): i.e., it
has been recorded from the Dzungarian Basin and Takla
Makan Desert in Xinjiang, China (A. Abdukadir in Zitt.
1993). The small southern populations in Baluchistan, isolated from the main population, occur in montane juniper
steppe (Roberts 1977). The manul’s range ends in thenorth where the steppes meet coniferous taiga forest(Bannikov 1954). It has been found at altitudes up to
4,800 m (Feng et ul. 1986), but it does not occur at suchhigh elevations as the snow leopard, and is more strongly
associated with flat, rolling steppe and south-facing slopes
where deep snow cover does not accumulate. Exposedrock outcrops or expanses of talus are a strong characteristic of its habitat (Heptner and Sludskii 1972). Manuls
have been collected from the fringes of cultivated areas in
China’s Qinghai province (Cai et al. 1989a).

Action Planning
Projects 77-80.

Population Status
Global: Category 4. Regional (Asia): Category 2. IUCN:
Insufficiently Known. The manul has been described as
most abundant on the cold grasslands of Mongolia andInner Mongolia (Mallon 1985, Feng et al. 1986, Y. Ma,
pers. comm. 1992). On the Tibetan Plateau, it occurs
widely but is nowhere common (G. Schaller in Zitt. 1993),
as most of the region lies above 4,500 m in elevation.
Elsewhere, the species is considered vulnerable to rare anduncommon: Afghanistan (Habibi 1977), Lakdakh, India(Mallon 1991), and Pakistan, especially the small, isolated populations found in Baluchistan (Roberts 1977).
In particular, the manul has disappeared in recent years
from much of the Caspian region (Bannikov and Sokolov
1984, Belousova 1993). Y. Ma (pers. comm. 1992) reportsthat it has been eliminated from the easternmost parts of its
range in China due to hunting.

Asiatic wildcat, Fe/issilvestris, ornata group(Gray, 1830)
Other Names
Asiatic desert wildcat, Asiatic steppe wildcat, Indian desertcat (English); chat sauvage d’Asie, chat orne (French);
Asiatische Wildkatze, Steppenkatze (German); gato montes, gato silvestre (Spanish); ye mao, caoyuan ban mao(Chinese); psk dsty (Dari: Afghanistan);velis cata
(Georgian); myallen, sabancha, myshuk dala, jawa misik(Kazakh); matsyl, zhapayi mishik (Kirgiz); jhang meno
(Kutch: India); tsookhondoi(Mongolia);
Asiaskiya
dkikaya stepnaya koshka, dlinahvostaya koshka, pyatnistaya koshka (Russian); yawa miishiik (Uygur); choi
pshak, sabancha, yobai pshak (Uzbek).

Protection
StatusCITES Appendix II. National legislation: lacking information. Hunting prohibited: Armenia, Azerbaijan, China,
India, Iran, Kazakhstan, Kyrgyzstan, Mongolia, Pakistan,
Russia, Turkmenistan,
Uzbekistan.
No information:
Afghanistan, Georgia, Tajikistan (Nichols et al. 1991,
Belousova 1993, IUCN Envl. Law Ctr. in Zitt. 1994).

Description
and Behavior (Plate 12)
The wildcats of central Asia differ from the European wildcats by having a more greyish-yellow
or reddish background color, marked distinctly with small black or
red-brown spots. The spots are sometimes fused into
stripes, especially in the central Asian regions east of the
Tian Shan Mountains (Groves 1980). The Asiatic wildcats
tend to be smaller in size, weighing between 3-4 kg(Schaller 1967, Roberts 1977), with females (mean 2.7 kg:

Principal Threats
Although there has been little recent international trade,the manul has long been hunted for its fur in relatively
large numbers. Western China’s annual harvest (exclud-99Part I: Species

Accounts.

Chapter

4. Eurasia,

Asiatic

wildcat

Litter size: (C) 2.75 (n=16; Mellen 1989); (W) up to 5-6(Ognev 1935, Sharma 1979).

Hemmer 1976) smaller than males. Some authorities consider the Asiatic wildcats and African wildcats to be conspecific (F. Zybica spp.), and the European wildcat (F.
silvestris) a separate species (Pocock 195 1, Ewer 1973,
Leyhausen 1979).
Like the other wildcats, rodents are the preferred prey:
jerboas, gerbils, voles, and mice (Ognev 1935, Allayarov
1963, Heptner and Sludskii 1972, Sharma 1979). The dietalso includes hares, young ungulates, birds, insects,
lizards, and snakes (Ognev 1935, Sapozhenkov 1961b,
Allayarov 1963, Lay 1967, Heptner and Sludskii 1972,
Roberts 1977, Sharma 1979). Sharma (1979) observed amother teaching her young to kill by bringing them injured
gerbils; she also provisioned them with beetles and eggs ofground birds. Asiatic wildcats rest and den in burrows(Ognev 1935, Allayarov 1963, Heptner and Sludskii 1972,
Sharma 1979). They are frequently observed in the daytime (Heptner and Sludskii 1972).

Age at sexual maturity: (W) 10 months (Roberts 1977), but
up to 21-22 months according to testicular development
in males (Heptner and Sludskii 1972).
Habitat and Distribution
Asiatic wildcats are most typically associated with scrub
desert (Allayarov 1963, Sharma 1979; T. Roberts in Zitt.
1993) (Fig. 4). They do not occur in the steppe grasslandsof Mongolia and Inner Mongolia (Zhang 199 1; X. Gao,
D. Mallon in Zitt. 1993), nor in alpine steppe (T. Roberts
in Zitt. 1993). They range up to 2,000-3,000 m in mountainareas with sufficient dense vegetation (Allayarov 1963,
Heptner and Sludskii 1972). Wildcats can be found near
cultivated areas (Salikhbaev 1950, Sharma 1979) andhuman settlement (T. Roberts in Zitt. 1993). They usuallyoccur in close proximity to water sources, but are also able
to live year-round in waterless desert. Snow depth limits
the northern boundaries of their range in winter (Heptnerand Sludskii 1972).
The Caucasus is the transitional zone between theEuropean wildcat (silvestris group) to the north and west,
and the Asiatic wildcat to the south and east. In thisregion, European wildcats are found in montane forest, andAsiatic wildcats are found in the low-lying desert andBiology
Reproductive season: (W) Mating season March-April andNovember-December(Rajasthan, India: Sharma andSankhala 1984); January-February (central Asia: Kashkarov 193 1, Allayarov 1963); year-round (Sind, Pakistan:
Roberts 1977).
Gestation: (C) 58-62 days (Hemmer 1976, Roberts 1977).

*:.-a*..-..

*

. .

.
‘.
*.
*.

:
.. .

-..

...*

.

.

.
-.

:

,. . .

..

.*

:
*.

*.*
Z,.’

-..

..

*..

*

Figure 4. Distributionof the Asiatic wildcat(E silwestris,
ornate group).
1. Khoshyeylag
I (Iran); 2. Registan Desert Wildlife Mgt. Reserve (proposed:
Afghanistan);
3. LalSuhanra V* (Pakistan);
4. Desert II; 5. National Chambal
IV; 6. Naroyan
SarovarIV; 7. Gir II complex(India); 8. Great Gobi II*; 9. Yolyn-am
I (Mongolia);
IO. Two specimens
collected from the Pune area,
India (Lamba 1967).

100

Part I: Species

Accounts.

Chapter

4. Eurasia,

Eurasianlynx

cats but less pale (Pocock 1939a, Kotwal 1984).
Roberts (1977) published reports of predation on
domestic poultry, but Heptner and Sludskii (1972) claimthat feral domestic cats and hybrids attack poultry more
often than wildcats.

semi-desert areas adjoining the Caspian Sea (Dal 1954,
Heptner and Sludskii 1972).
Population Status
Global: Category 5c. Regional (Asia): Category 2. IUCN:
not listed. In the central part of its range, Belousova(1993) and E. Matjuschkin (in Zitt. 1993) report that the
wildcat is common and populations stable in the lowlandsof Kazakhstan. In Azerbaijan, the ornata-silvestristransition zone, a pronounced loss of range has been documented (Belousova 1993). In India, the eastern limit of
its range, the Wildlife Institute of India (in Zitt. 1992) considers that 90% of the species’ habitat in India has beenlost. On the other hand, Sharma (1979), who studied thespecies in western Rajasthan, noted that the introduced
mesquite Prosopis juliflora, which provides favorable
habitat for the wildcat, was spreading extensively in various regions of the Indian desert.

Action Planning
Projects 10, 15, and 80.

Eurasian lynx, Lynx lynx
(Linnaeus, 1758)
Other Names
Lynx (French); Luchs (German); lince (Spanish); sinokoi(Ainu: Sakhalin island); lusan (Armenia); meshag, mesh
(Azerbaijan);tsogde (Baltistan:
Pakistan);shihli(Chinese); ilves (Finland); lynx (French); potskhveri(Georgia); varchakh (Farsi: Iran); lince (Italy); patsalam(Kashmiri); silovsin, suloosun (Kazakh; Kyrgyz, Uzbek);yi (Ladakhi: India); phiauku (Lahul: India); shleleisin(Mongolia);
gaupe (Norway);rys (Russian: Czech
Republic, Slovakia, Romania, Russia, Slovenia); lodjur(Sweden); vasak (Turkey); su laisun (Uygur).

Protection
StatusCITES Appendix II. National legislation: fully protectedin the east of its range; elsewhere hunted commercially or
not protected.
Hunting and trade prohibited: India,
Pakistan. Hunting and trade regulated: China, Kazakhstan,
Kyrgyzstan, Turkmenistan, Tajikistan, Uzbekistan. No
legal protection: Georgia, Iran, Mongolia. No information: Armenia, Azerbaijan (Nichols et al. 199 1, Belousova
1993, A. Bukhnicashvili
in Zitt. 1993, IUCN Envl. Law
Ctr. in Zitt. 1994).

Description
and Behavior (Plate 11)
The Eurasian lynx is the largest of the lynxes. Adult males
weigh on average 2 1.6 kg (n= 103), while females areslightly smaller at 18.1 kg (n=93). The lynxes of eastern
Siberia consistently reach the greatest size (Breitenmoser
and Breitenmoser-Wursten
in prep.). The Eurasian lynx
has relatively long legs, and large feet which provide a“snowshoe effect,” allowing for more efficient travel
through deep snow. In winter, the fur grows very densely
on the bottom of the feet (Formozov 1946). The coat is
greyish, with tint varying from rusty to yellowish.
Abright reddish tint, with profuse spotting, is seen most frequently in the southwestern part of the lynx’s range (southern Europe, Asia Minor and the Caucasus: Heptner andSludskii 1972).
There are three main coat patterns: predominantly spotted, predominantly striped, and unpatterned. While thespotted-striped types, controlled by the “Tabby” gene, predominate in present reintroduced European lynx populations (originating mainly from the Carpathian mountains
further east), Ragni et al. (1993) show through examination of 26 pelts of the original, now extinct, populationsof the European Alps that these animals were chiefly
unpatterned, and were, moreover, smaller in size.
Eurasian lynx have long, prominent black ear tufts, andshort black-tipped tails. Lynx activity peaks in the evening
and morning hours, with resting mainly around mid-day
and midnight (Bemhart 1990).

Principal Threats
In the past, Asiatic wildcats have been trapped in large
numbers in several areas: e.g., 12,800 in Kazahkstan(19289: Ognev 1935); 1,350 in the Kyzylkum desert(Allayarov 1963); 1,500 annually in the 1980s in Xinjiang(X.-Y. Gao in Zitt. 1992). In 1979, traders in India
declared stocks of 41,845 pelts for an export amnesty(Panwar and Gopal 1984). Habibi (1977) reports widespread hunting of the wildcat for the fur trade in
Afghanistan, and that large numbers of pelts were seen forsale in Kabul bazaars. Roberts (1977) equates the cat’s
rarity in Pakistan with demand from the fur trade.
However, at present there is little international trade in
Asiatic wildcats (WCMC unpubl. data).
Hybridization with domestic cats has been reported
from Pakistan (Roberts 1977 and in Zitt. 1993) and central
Asia. Heptner and Sludskii (1972: 491) state that “thefemale [Asiatic wildcat] mates quite often with a domestic
male, and hybrid offspring are frequently found near villages where wild females live.” The situation in other partsof its range, including India, is probably similar. It has
been reported that the most common race of the domestic
cat occurring in rural areas in India is colored dark grey,with black stripes and spots, similar in appearance to wild-101

Pat-t I: Species

Accounts.

Chapter

4. Eurasia,

Eurasianlynx

Although the Eurasian lynx is often classified with thethree other lynxes as a predator of lagomorphs (e.g.,
Gittleman 1985), this is a major misconception (Breitenmoser and Breitenmoser-Wiirsten
in prep.). Small ungulates, particularly roe deer, chamois, and musk deer, arethe main prey, and lynx will generally only take small prey
when ungulates are scarce (Vasiliu and Decei 1964,
Danilov et al. 1979, Birkeland and Myrberget 1980,
Heptner and Sludskij 1980, Jonsson 1980, Somerlatte et al.
1980, Pulliainen 198 1, Malafeev and Kryazhimsky 1984,
Breitenmoser and Haller 1987, Herrenschmidt and Leger
1987, Dunker 1988, Hucht-Ciorga 1988, Cop 1992, Ragni

Species.

range

Zone of greatest lynx abundancein Russia according
to Matjuschkincl

et al. 1992, Zheltuchin 1992). When young blue sheep
are not available (A. Abkukadir, pers. comm. 1992), lynx
in China have been reported to prey on pikas, large
rodents, and hares (Feng et al. 1986). Pulliainen et al.
(1988) point out that, in Finland, lynx tend to be in better
condition in the southwest-where
there is an introduced
population of white-tailed deer from North America-thanin the remainder of the country, where roe deer are veryrare and hares are the main available prey. Similarly,
Zheltuchin (1992) states that lynx are found at lower densities in the northern parts of Siberia where there are haresbut no ungulates; in these regions, arctic hares and lynx

Protected area where
species occurs
Protected

area where(1978)

Figure 5. Distributionof the Eurasianlynx (L. lynx).
1. Bialowieza
II# (Poland) + Belovezhskaya
Pushcha IV (Byelorus)
complex;
2. Munzur
II (Turkey);
3. Borzhom
I (Georgia,
possibly now extirpated:
Z. Gurielidze
and A. Bukhnicashviliin litt. 1993); 4. Lisar V;
5. Alborz-e-Markazi
V complex (Iran); .6 Pamir-i-Buzurg
IV; 7. Ajar Valley IV (Afghanistan);
8. Khunjerab
II (Pakistan);
9. Hemis II (India);
10. Taxkorgan
IV; 11. Qomolongma
IV; 12 Arjin Mts. IV complex;
13. Wolong IV*; 14. Boghdad
Mts. IX*; 15. Hanas IV; 16 Changbai
Mts.
IV* (China); 17. Great Gobi II* (Mongolia);
18. Pechoro-Ilych
I; 19. Stolby I (Russia).

102

Part I: Species

Chapter

4. Eurasia,

Eurasianlynx

Age at last reproduction: (W) females 14 years (n=l);males 16-17 years (n=l: U. Breitenmoser in Zitt. 1993).

fluctuate cyclically (Heptner and Sludskii 1972), similarly
to fluctuations described for the Canada lynx (see Species
Account).
In some parts of their range, lynx prey mainly on large
ungulate species (mostly females or young), including red
deer (Hell 1973, Gossow and Honsig-Erlenburg
1986,
Jedrzejewski
et al. 1993), reindeer (Haglund 1966,
Bjarvall 1992), and argali (Matjuschkin 1978). Lynx arecapable of killing prey 3-4 times their own size (Gossow
and Honsig-Erlenburg 1986, Haller 1992).
While all the lynx species are similar in appearance, theEurasian lynx bears the closest morphological resemblance to the Canada lynx (Kurten and Rausch 1959), andthe two are very often treated as conspecific. However,
Breitenmoser and Breitenmoser-Wiirsten(in prep.) argue
convincingly that the two are ecologically separate species.
Specialization for different prey has led to a divergence inlife history and social and spatial organization. Unlike
the Canada lynx, the Eurasian lynx has a “phenotype set”
typical of a large felid (Sunquist and Sunquist 1989): it islarge, long-lived, kills prey at least half its own bodyweight, forages over wide areas, and generally exists at
low densities. Only in some parts of its range, chiefly thenorthern boundary, is there ecological similarity between
the Eurasian and Canada lynxes in their predation on cyclically fluctuating hare populations.

Longevity: (W) up to 17 years (Kvam 1990, U. Breitenmoser in Zitt. 1993); (C) up to 24 years (Green 1991).
Habitat and Distribution
Throughout Europe and Siberia, lynx are associated primarily with forested areas which have good ungulate populations (Haglund 1966, Novkov and Hanzl 1968,
Matjuschkin 1978, Malafeev and Kryazhimskiy
1984,
Haller and Breitenmoser 1986, Breitenmoser and Haller
1987). In central Asia, lynx occur in more open, thinly
wooded areas (Heptner and Sludskii 1972, Matjushkin
1978, Tan 1984). Lynx are probably found throughout
the northern slopes of the Himalayas, and have beenreported both from thick scrub woodland (Chundawat
1990a) and from barren, rocky areas above the treeline(Roberts 1977). On the better-forested southern Himalayan slopes, the only record is a sighting in alpine tundra(4,500 m) from the Dhaulagiri region of Nepal (Fox 1985,
D. Mallon in Zitt. 199 1). Lynx occur locally over theentire Tibetan plateau, and are found throughout the rocky
hills and mountains of the central Asian desert regions(Bannikov 1954, Stubbe and Chotolchu 1968, Heptnerand Sludskii 1972).
The Eurasian lynx has one of the widest ranges of all
cat species, with approximately 75% of the range within
the borders of Russia (Fig. 5). Lynx have been recorded as
far north as 72” N, near the edge of the continental landmass (Zheltuchin 1992).

Biology
Reproductive season: (W) mating season February-April,births May-June (Europe, Russia: Ognev 1935, Dal 1954,
Kazcensky 1991, Kvam 1991).

Population Status
Global: Category 5b. Regional (Asia): Category 3.
Regional (Europe): Category 2. IUCN: not listed. The
stronghold of the Eurasian lynx is a broad strip of southem Siberian woodland stretching through Russia from theUral mountains to the Pacific. The Russian population
has been estimated to be 36,000-40,000 (Matjuschkin
1978, Zheltuchin 1992), but it is not clear how these figures
were derived (U. Breitenmoser, pers. comm.). Heptner andSludskii (1972) reviewed reports on lynx distribution indetail, and concluded that in Russia a major population
increase and range expansion (including the colonizationof the entire Kamchatka peninsula) took place in the 193Os1940s. Lynx re-colonized areas where they had previously
been extirpated, mainly due to a sharp decline in commercial hunting during this period of social upheaval.
In China, lynx are found throughout much of the country, concentrated in the montane regions. Given its wide
distribution, Tan Bangjie (in Zitt. 1987) and A. Abdukadir(in Zitt. 1993) are relatively optimistic, but emphasize thatin many places it has become locally rare. Ma Yiqing

Gestation: (C) average 69 days (Hemmer 1976).
Litter size: (W) 2.5 k 0.5 (Norway, n=8: Kvam 1991); 1.82
& 0.6 (Switzerland, n=14: U. Breitenmoser in Zitt. 1993);(C) 2.1 k 0.9; range l-4 (n=141: Kaczensky 1991).
Interbirth interval: (W) generally one year, but with occasional breaks, e.g., three years with litters, one without(Switzerland: U. Breitenmoser in Zitt. 1993).
Age at independence: (W)Breitenmoser et al. 1993a).

Accounts.

10 months (Switzerland:

Age at first reproduction: (W) females 20-24 months(Kvam 199 1, U. Breitenmoser in Zitt. 1993); males approximately 30 months (Kvam 199 1).
Juvenile mortality: (W) Breitenmoser et al. (1993a) foundhigh rates of juvenile mortality for a lynx population living
in a densely settled area of Switzerland: 50% pre-dispersal (n= 14 kittens); 80% post-dispersal (n=5 sub-adults).
Recruitment rates: (W) 0.69 (Breitenmoser et al. 1993a) 1.25 k 1.5 (Kvam 1990) juvenile lynx per female per year.

103Part I: Species

Accounts.

Chapter

4. Eurasia,

Eurasianlynx

with the spotted Iberian lynx, L. pardinus (Heptner andSludskii 1972).
The most comprehensive data on species status is fromthe European sub-region (Breitenmoser and BreitenmoserWiirsten 1990, Anon. 1992b), where lynx are thinly distributed and isolated into discrete sub-populations (Fig. 6).
The species was actually eradicated from most of the subregion within the past 150 years (Kratochvil 1968), surviving only in the north and the east. In these regions,
numbers fell in the early 1900s but recovered concurrently
with increases in small ungulate populations (Breitenmoser and Breitenmoser-Wursten
1990). Lynx have since
been reintroduced in several parts of western Europe, themost extensive effort for any felid species (see Part II
Chapter 6).
In northern Europe (Finland, Norway, Sweden, northeastern Poland, and the European region of the former
U.S.S.R.), the population is stable and connected to thelarger Siberian population. In central Europe, a relatively
large but isolated population is found in the Carpathian
Mountains (Slovakia, Poland, Romania, Ukraine). Small(pers. comm. 1992) believes populations are declining in
the northeast. G. Schaller (in Zitt. 1993) notes that lynx
are the most commonly seen cat pelts in local fur markets
in the west.
Little information is available from the remainder of thelynx’s wide Asian range. In Ladakh, Mallon’s (1991) survey indicates that it is rare in the central region, but
Chundawat (1990a) found it locally common in dense
thicket scrub in the Nubra river valley. The lynxes of thecentral Asian deserts and high mountains inhabit ecosystems very different from the cold coniferous forests withwhich the species is primarily associated. They appear toprey mainly on hares and rodents, rather than ungulates,
but their ecology is little known (A. Abdukadir in Zitt.
1993, U. Breitenmoser, pers. comm.). Bannikov (1954)
described lynx as common in the desert hills of southwestern Mongolia. Lynx are now quite rare in the Caucasus (Z.
Gurielidze and A. Bukhnicashvili in Z&t. 1993). Animals
from this region, with their small size, reddish coat, andheavy spotting, are sometimes recognized as the subspecies L.Z. din&i, and were once considered conspecificFigure 6. Recent distributionof Lynx lynx in Europe (Breitenmoser
and Breitenmoser-Wiirsten
1990).
Dark shading: occupied area. Light shading: occasionallyoccupied area, or area with low
population
density according
to survey respondents.
Stippling:
lynx range according
to the literature.
Dotted line: northern and southernboundaries
of lynx range in the former U.S.S.R. (Matjuschkin1978); see Figure 5. Asterisk (*): isolated observations.
(AL = Albania, AT = Austria, BG = Bulgaria,
CH = Switzerland,
CS = Czech Republic + Slovakia,
DE+DD = Germany,
ES = Spain, FR = France,
GR = Greece, HU = Hungary,
IT = Italy, NO = Norway,
PL = Poland, RO = Romania,
SE = Sweden,
SF = Finland, SU = former U.S.S.R., TR = Turkey, YU = former Yugoslavia.

104Part I: Species

populations are found in the French Pyrenees and Vosges
mountains; the Jura Mountains (France, Switzerland); theAlps (Austria, France, Italy, Switzerland); the Balkans(Albania, Croatia and Slovenia); and the Bohemian forest(Czech Republic) (Breitenmoser 199 1).
The most thorough estimates of resident adult density(per 100 kmz), derived from radiotelemetry studies, areavailable from Switzerland:
0.94 (Jura Mountains:
Breitenmoser et ul. 1993a); 1.2 (northern Alps: Haller andBreitenmoser 1986); 1.43 (central Alps: Haller 1992).
Based on snow tracking, Hjelm (199 1) estimated 0.340.74 individuals/l00
km2 in Sweden. Where ungulate
prey is abundant, density estimates are high: lo- 19 lynx/
100 km2 in the Bialowieza Forest in Poland and Byelorus(Heptner and Sludskii 1972).
Where hares are the major prey, density estimates from
Russia are of the order of less than four lynx per 100 km2(Zheltuchin 1992).
Excluding outliers, Breitenmoser
et al. (1993a)reported average home ranges for males of 264 t 23 km2,
and 168 k 64 km2 for females. Within these home ranges,
core areas averaged 185 t 58 km2 for males, and 72 k 27
km2 for females. Females tended to use the central partof their home ranges more intensively (Kaczensky 199 l),
whereas males regularly visited the periphery of theirhome ranges (Diitterer 1992). Thus, male core areas averaged 70% of their home ranges and showed some overlap, while those of females were exclusive, and averaged
only 44% of their home ranges. With the exception of theoverlap zones, one male and one female shared the same
area. On average, 86% of a female’s home range was covered by a male’s home range. Studies from Sweden (Haglund 1966) and Russia (Matjuschkin
1978, Zheltuchin1984) have also concluded that males generally share theirranges with just one female and her kittens. However,males seem to avoid female core areas, and thus appear tocontrol a zone around females and their kittens, avoiding
competition for prey and excluding other male competitors(Breitenmoser et al. 1993a).

Accounts.

Chapter

4. Eurasia,

Eurasianlynx

Principal Threats
Lynx are vulnerable to destruction of their ungulate preybase. Under harsh winter conditions, they may not be able
to subsist successfully on smaller prey (Pulliainen 1992).
Large ungulate prey are favored in the winter because of
their vulnerability in deep snow. For example, Scandinavian lynxes have been reported to switch from predation
on small game in autumn to large game in winter(Haglund 1966, Birkeland and Myrberget 1980). Huntingpressure may also play a role in lynx population declines(Hell 1992).
Zheltuchin (1992) reported that clear-cutting can have anegative effect on lynx abundance. In the Tver region,
lynx were stable and resident when the level of deforestation was approximately 25%. When 80% of an area wasclear-cut, the frequency of lynx tracks was about 15 times
lower than in areas consisting of 40-50% mature forest
cover.
Breitenmoser
and Breitenmoser-Wursten(1990)
review lynx predation on livestock for European countries,
and include information on the different ways it is dealt
with by national authorities (see also Part II, Chapter 2).
Problems are most severe in western Europe where lynx
have been reintroduced. After native wild ungulates readapted to the presence of predators, livestock killingincreased, but later declined as lynx dispersed and becameless concentrated. Overall stock losses are relatively low
in these countries, and are compensated either by the government or environmental groups. Switzerland, whichinvests about U.S. $35 million every year as a subsidy forsheep farming, pays out only about U.S. $7,000 (Anon.
1994a) as annual compensation for lynx kills (rates areagreed upon by stock owners). The problem is thus notreally economic, but psychological and political (Breitenmoser and Breitenmoser-Wiirsten
1990). For 100 years,
western European farmers have had the luxury of not having to guard livestock against losses to predators. A
prominent French farmer invited to speak at a symposiumon the lynx referred to it as “a savage and outdated animal”(Grosjean 1992).
There is no information beyond harvest reports on
which to base an assessment of the biological impact ofcommercial trapping for furs, and thus its significance asa threat is difficult to judge. In Russia, A. Zheltuchin(1992, and in Breitenmoser and Breitenmoser-Wursten
1990) and Matjuschkin (1978) indicated that harvest levels
range between 2,000-5,800 annually. The maximum harvest reported (5,800 in 1956) is similar to harvests reported
for 1985- 1986, which could indicate that the lynx population has remained relatively stable. Annual harvests on theorder of 5,000-6,000 have been reported in the Soviet
Union as long ago as 1928 (Heptner and Sludskii 1972).
Russia exports most of its reported harvest, averaging
about 5,000 pelts per year between 1985- 1989 (WCMC

Protection
StatusCITES Appendix II. Hunting prohibited: Albania, Austria,
Bulgaria, Czech Republic, France, Georgia, Germany,
Greece, Hungary, India, Iran, Kazakhstan, Kyrgyzstan,
Nepal, Pakistan, Switzerland, Tajikistan, Turkmenistan,
Uzbekistan. Hunting regulated: China, Finland, Slovakia,
Mongolia, Norway, Poland, Romania, Sweden, Russia,
Turkey.
Hunting prohibited in protected areas only:
Bhutan, Myanmar. No information: Afghanistan, Armenia, Azerbaijan, Belarus, Estonia, Iraq, Italy, North Korea,
Latvia, Lichtenstein, Lithuania, Slovenia, Syria (Breitenmoser and Breitenmoser-Wursten
1990, Nichols et al.
1991, Anon. 1992b; A. Bukhnicashvili, E. Mukhina in litt.
1993, IUCN Envl. Law Ctr. in litt. 1994).

105

Part I: Species

Accounts.

Chapter

4. Eurasia,

lberianlynx

unpubl. data). There was a surge in exports from China
from 1984- 1988, with a peak of over 12,000 skins in 1986
(WCMC unpubl. data). This trade was probably inresponse to high pelt prices prevailing at that time, when
Canada lynx populations were at a cyclic low. Given that
China lacks the organized trapping infrastructure present
in Russia, it is possible that the skins could have beentaken originally in Siberia, unaccounted for in the official
harvest (U. Breitenmoser in Zitt. 1992).
Both China and Russia announced in 1993 the settingof export quotas for lynx furs: 2,800 per year from Russia,
and 1,000 per year from China (Anon. 1993b). Exportsof lynx furs from these two countries are currently low,
below 1,000 annually.
While lvnx reintroduction in Switzerland has been considered a success, Breitenmoser et al. (1994) have foundthat the population has stopped expanding, and is threatened by an imbalanced sex ratio (lack of males). The problems facing the Swiss lynx population are discussed in Part
II, Chapters 2, 3, and 6. In general, lynx adapt well to settled and cultivated areas if population levels do not
become too low. Lynx have been reported from the outskirts of Moscow, Leningrad, and other large Russian
towns (Heptner and Sludskii 1972).

the Eurasian and Iberian lynxes evolved from the first
identifiable lynx, Lynx issiodorensis-theIberian inEurope, and the Eurasian lynx (which gave rise to the
Canada lynx) in China. Although the ranges of theEurasian and Iberian lynx never overlapped very much,and have become essentially separate in recent times, thetwo lynxes may still co-exist in the Pyrenees Mountains
between France and Spain (van den Brink 197 I,
Breitenmoser and Breitenmoser-Wiirsten
1990).
The ecology of the Iberian lynx is very different fromthe Eurasian lynx. While the Eurasian lynx is a forest animal which preys on ungulates, the Iberian lynx is found in
scrub vegetation and preys almost exclusively on European
rabbits. In both ecology and average body weight, theIberian lynx is very similar to the Canada lynx and bobcat
of North America. By weight, 93% of lynx prey during thesummer season is made up of rabbits, which suffer particularly at that time from the poxvirus myxomatosis. The
proportion of rabbits in the diet decreases slightly in thewinter months, when rabbit numbers are at an annual low(Delibes 1980, Beltran et al. 1987). At this time, red deer
(fawns), fallow deer, and moufflon (juveniles) are taken(Aymerich 1982, Beltran et al. 1985). In the Coto Dofiana
wetland area along the southwestern Spanish coast, ducks
are a seasonally important food resource from March toMay, during their breeding season (Delibes 1980, Beltranand Delibes 1991). The energy requirements of the Iberian
lynx have been estimated at approximately one rabbit per
day (Aldama et al. 1991).
A radio-telemetry study in the Coto Donana National
Park showed lynxes to be primarily nocturnal, with activity peaking at twilight as the animals moved out of their
daytime resting places to hunt. Daily travel distance averaged 7 km, with males generally travelling further thanfemales. Diurnal activity peaks during the winter (Beltran
et al. 1987).

Action Planning
Projects 16, 80, and 84-88.

Europe Sub-region
Iberian lynx, Lynx pardinus(Temminck, 1827)
Other Names
Pardel lynx, Spanish lynx (English); lynx d’Espagne(French); Pardelluchs (German); lince iberico (Spanish);lobo cerval (Portugese).

BiologyMating season: (W) January-July, peak January-February.
Birth season: (W) March-April

peak.

Gestation: (W) approx. two months.

Description
and Behavior (Plate 11)
The Iberian lynx looks like a smaller version of theEurasian lynx, being only about half its size, with adultmales weighing an average of 12.8 kg (n=5) and females
9.3 kg (n=4) (Beltran and Delibes 1993). Iberian lynxes
have a distinctly spotted coat, as do Eurasian lynxes ofwestern Europe. However, the two are different species(Werdelin 1990, Garcia-Perea 1992), sympatric in central
Europe during the Pleistocene (Kurten 1968, Kurten andGrandqvist 1987), with the time of separation estimatedto have occurred long before the separation of the Eurasianand Canadian lynxes. Werdelin (198 1) considers that both

Litter size: (W) 2-3 (M. Delibes in Zitt. 1993).
Survival to independence: (W) l-2 kittens per female.
Age at independence: (W) 7-10 months.
Age at dispersal: (W) independent kittens remain in theirnatal territory until an average of 20 months (range 8-28;n=15).
Age atfirst reproduction: (W) Females are able to breed intheir first winter, but the time of first reproduction depends
upon demographic and environmental factors. In a high-106

Pat? I: Species

Accounts.

Chapter

4. Eurasia,

lberianlynx

heather scrub (Beltran et al. 1987).
The Iberian lynx has historically been restricted to the
Iberian peninsula, where it was widespread (Graells 1897),
and southern France (Lavauden 1930). The peninsula wasapparently a Pleistocene refuge for the European rabbit, andtoday the race that occupies this area is only half the sizeof conspecifics found elsewhere in central Europe (1 vs. 2
kg: Gibb 1990). The Iberian peninsula is the only part of
the Palearctic region which supports a relatively high density of lagomorphs, similar to that found in North America,
home to two species of lagomorph-eating lynxes: the bobcat and Canada lynx (U. Breitenmoser, pers. comm. 1992).
By the early years of the 20th century, the Iberian lynx
had become very rare in northern Spain, although it was
still abundant in the center and south (Cabrera 19 14). By
the 1960s its range was essentially limited to the southwestern quarter of the peninsula, an area of some 57,000km”, where the population probably had a continuous distribution (Rodriguez and Delibes 1990). At present, lynx
range in Spain (where 95% of the population is now
found) covers only 14,000 km2, of which about 11,000
km2 is believed to be breeding range. This represents only
about 2% of the country’s total area (Rodriguez anddensity population, such as that in Dofiana NP, age at first
reproduction depends upon when a female acquires a territory. This normally occurs because of either death or
expulsion of a resident. One female did not reproduce
until five years of age, and this only occurred when themother died and left the territory vacant (J. Aldama, P.
Ferreras in Zitt. 1993).
Age at last reproduction: (W) 10 years (male and female:
M. Delibes in Zitt. 1993).
Longevity: (W) up to 13 years (Ferreras et al. 1992).
Habitat and Distribution
The Iberian lynx occurs in Mediterranean woodland andmaquis thicket. It favors a mosaic of dense scrub for shelter and open pasture for hunting rabbits (ICONA 1992).
Palomares et al. (199 1) examined habitat preferences oflynx in the Coto Donana area of southwestern Spain,
including the national park and environs. Lynx were generally absent from cropland and exotic tree plantations(eucalyptus and pine), where rabbits were also scarce. In
the park, radiotelemetry showed that more than 90% of
daytime resting spots used by lynx were located in thicka

I 11 IKm

Figure 7. Distributionof the lberian lynx (L. pardinus)
after Rodriguez
and Delibes (1992).
1. Serra da Malcata IV (Portugal);
2. Monfrague
V; 3. Cabafieros
V; 4. Sierra de Aracena y Picas deAroche V; 5. Sierra Norte Natural Park; 6. Sierra de Hornachuelos
V; 7. Sierra de Carde:a y Montoro;
8. Sierra de Andujar V; 9. Despefiaperros
V; 10. Cazoria, Segura y la Villas Natural Park; 11. Sierra
Mgina V; 12. Entorno de Dofiana V; 13. Dofiana II* (Spain).

107

Par? I: Species

Accounts.

Chapter

4. Eurasia,

lberianlynx

Population Status
Global: Category 1. Regional (Europe): Category 1.
IUCN: Endangered. The Iberian lynx is the only cat
species ranked in Category 1. The total number of Iberianlynx, including sub-adults but not kittens, probably does
not exceed 1,200, with only about 350 breeding females(ICONA 1992, Rodriguez and Delibes 1992). The lynx
population is extremely fragmented. In Spain, a comprehensive survey (Rodriguez and Delibes 1992) documented
48 isolated breeding areas, 32 areas of occasional presence, and 50 other areas where lynx presence is suspected
but not confirmed (Fig. 7). Since lynx are known to dis-

Delibes 1992). Distribution in Portugal is less wellknown, but has also been substantially reduced since the1940s. There now appear to be only three breeding subpopulations in that country, occupying a total range of only
about 700 km2, with the largest now found in the Serra da
Malcata Nature Reserve and the Algarve Mountains of
the extreme south (Delibes 1979, Palma 1980, ICONA
1992). Lynx distribution is centered on mountain ranges,
where land use is mainly in the form of privately owned
hunting reserves (ICONA 1992). Lynx are mainly foundbetween 400-900 m elevation, but will range up to 1,600 m(IUCN 19762, Palma 1980).

N

GAT

t

CP

=igure 8. Population
structureof the lberian lynx in Spain (Rodriguez
and Delibes 1992).
Zontinous lines surroundthe nine estimatedpopulations,
including breeding (shaded black) and occasional presence areas (stippled).
The breeding populations
are the same as those shown in Figure 7.
Broken lines further delineate three sub-populations(VIL, MTO, SMO).
Straight lines represent
minimum barrier breadth (km). Symbols indicate the degree of barrier penetrability
for lynx: star = high;solid circle = low; open circle = null. GAT = Sierra de Gata. GRE = Sierra de Gredos.
MAD = Alto
Alberche.
SSP = Sierra de San Pedro. CP = Central population(VIL = Villuercas;
MT0 = Montes deToledo; SMO = Eastern Sierra Morena).
SMH = Western Sierra Morena. SMC = Central Sierra
Morena.
DON = Donana; SUB = Sierra Subbeticas.

108

Par? I: Species

Table 3Comparative
ImportanceAmong LyrWS
in Spain(%) of Different

Accounts.

Chapter

4. Eurasia,

lberianlynx

Causes of Mortality(after Rodriguez

and Delibes

1990)
Other

N

8.2

170

0.1

8.6

689

6.7

7.0

15.7

356

4.0

2.1

10.6

1,215

PeriodGuns

Traps/
Snares

Dogs

1958

21.2

67.0

3.5

1958-1977

26.0

62.7

2.6

1978-1988

26.1

44.4

Total

25.4

58.0

Road

1950s;the lynx wasnot declareda protected speciesuntil
1973 (ICONA 1992). At present, the fine for willful
killing of a lynx is very high, approximately U.S. $8,000(Delibes 1989).

perseup to 30 km (Beltran 1988), the 48 isolated breeding areasand 32 areasof occasionalpresenceare likely to
makeup nine distinct sub-populations(Fig. 8). Thesesubpopulationsareprobably genetically isolated,having been
separatedby intensive agriculture and settlement by anaverage distance of 45 km. Only two sub-populations
occupy areaslarger than 2,000 km”.
Only the central population, consisting of three subpopulations(VIL, MT0 & SMO on Fig. S), is believed to
be viable, consisting of some 800 lynx. The remaining
sub-populationsare estimatedat between 13-63 animals(Rodriguez and Delibes 1992). Small population size is aproven threat to the Iberian lynx: it hasdisappearedfrom
91% of the areaslessthan 1,000 km2 in size which were
estimated to have harbored it in 1960 (Rodriguez andDelibes 1990).
Iberian lynxes have beenstudied usingradiotelemetry
in the Coto DofianaNP since 1983. In good quality habitat, lynx density (including sub-adultsbut not kittens) was
estimatedat 16 individuals per 100 km2 (Palomareset al.
1991). Rodriguez and Delibes( 1992)estimateddensities
acrosslynx rangebasedon the relationshipbetweenreportsof lynx presenceand actual numberspresent,known fromthe Dofiana study area. Densitiesfor the nine geneticallyisolated sub-populationswere estimatedat between 4.510.1 individuals per 100 km? For resident adults in the
Dofiana, annual home range averages 18 km2 for males(monthly home range averages 10 km2) and 10 km2 forfemales(monthly homerangeaverages8 km2) (M. Delibes
in Zitr. 1993). Home ranges are intrasexually exclusive,with completeintersexualoverlap (Beltran et al. 1987).

Principal Threats
The decline of the lynx population since the 1960shas
beenprimarily causedby habitat lossanda declineof theirmain prey species,the European rabbit. The poxvirus,
myxomatosis,was introduced from South America in theearly 1950sandhada devastatingimpacton Europeanrabbits, which had no natural immunity. In the early yearsof
the epidemic, rabbits virtually disappearedfrom many
areas. Europeanrabbits are in the processof developing
geneticresistanceto myxomatosis,which is no longer suchan important threat. However, a new disease,viral hemmorrhagicpneumonia,hit the Spanishpopulation in 1988,
causinghigh initial mortality of adult rabbits (Gibb 1990,
Villafuerte and Moreno 1991). At the sametime, largescale habitat conversion has taken place in Spain andPortugal, where the pasture-scrub-woodlandmosaicpreferred by rabbits was replacedby wheat fields and industrial forest plantations. Rabbits are declining even in themontanehunting reserves,probably becausesmall-scalegrazing and cultivation have been abandonedin theseareas,and the pasturelandpreferred by rabbits is invaded
by thicket (ICONA 1992).
Nevertheless,there are someareaswhere habitat quality and rabbit density appear sufficient, yet no lynx are
found. Particularly in theseareas,it seemsthat humansare
directly responsiblefor an appreciablelevel of lynx mortality (Delibes 1989). This is true even for the population
living in the area receiving the greatest protection, theDofiana NP complex. Most of the deathsrecorded there
in the last 10 yearswere human-related,and only 8.3% of
the annualmortality rate can be related unequivocally toProtection Status
CITES Appendix I. National legislation:fully protectedin
Spain and Portugal (ICONA 1992). The Spanishgovernment paid a bounty for destruction of lynx up through the

IO9Part I: Species

Accounts.

Chapter

4. Eurasia,

Europeanwildcatnatural causes (Ferreras et al. 1992). Rodriguez andDelibes (1990) compiled records on cause of death for
1,2 15 lynx killed in Spain over the past 30 years.
Traps and snares, particularly gin traps set for rabbits,
have been the principal known cause of death for lynx,
although the practice of trapping rabbits is now declining.
Road deaths were comparatively unimportant (or seldom
reported) before 1978, but are expected to increase as
Spain undertakes an ambitious program of road-buildingin the 1990s (ICONA 1992).
The small, isolated sub-populations of Iberian lynx aretheoretically vulnerable to genetic drift, where alleles withlow frequency are likely to disappear from the population
gene pool. Beltran and Delibes (1993) found preliminary
evidence for this happening in Coto Dofiana, where thepopulation of approximately 40-50 lynx has been isolated
since the early 1960s. Three pelage patterns were present
in the population at that time, but now no animals exhibit
the rarer fine-spotted pattern.
The Spanish government is in the process of developinga national conservation strategy for the Iberian lynx, withthe goal of enabling the lynx to occupy as large a range as
possible on a permanent basis. Management measures
will be applied first to the largest population nuclei (theeastern Sierra Morena, the Toledo Mountains, the corridors between these two zones, and certain parts ofExtremadura). Measures include completion of detailed
surveys of the conditions faced by each lynx sub-population (land use, land ownership, habitat condition, rabbit
density); banning rabbit trapping; taking active steps toincrease rabbit populations (such as brush clearance); andestablishment of a captive breeding program (now underway) (Rodriguez and Delibes 1990, ICONA 1992).

(Slovakian);
yaban kedisi(Wallon: Belgium).

sauvadge tche

Description
and Behavior (Plate 12)
The forest wildcats of Europe and western Russia are greybrown in coat color, with bushy, blunt-ended tails and awell-defined pattern of black stripes. Although they tend
to look bigger than African wildcats because of their thick
winter fur, an extensive series of weight measurements
have shown that they are not: males weigh an average of
5 kg and females 3.5 kg (Conde and Schauenberg 197 1).
However, the authors did record strong seasonal weightfluctuations ranging up to 2.5 kg, with heaviest maleweights recorded from September to the end of February(France).
The fossil record suggests that the European form of
the wildcat is the oldest, descended from Martelli’s cat(Felis [silvestris]
lunensis) about 250,000 years ago(Kurten 1968). Molecular analysis indicates that theAfrican wildcat diverged from the European form only
about 20,000 years ago (Randi and Ragni 1991). This iscorroborated by the fact that fossil specimens of African
wildcats are only known with certainty from the late
Pleistocene (Savage 1978). The domestic cat was derived
from African wildcats between 4,000-8,000 years ago(Clutton-Brock
198 1, Davis 1987, Kitchener 1992).
Hybridization is common between European wildcats anddomestic cats, and Kitchener (1992) discusses characters(pelage pattern, gut length, skull morphology) that can beused to distinguish reliably pure wildcats from hybrids ordomestic tabbies. Many hybrids are more like wildcats
in size and morphology than domestic cats: perhaps there
is differential survival of hybrid forms in the wild thatfavors larger cats. Large black cats observed in Scotland(“Kellas cats”) and the Caucasus (Satunin 1904, Aliev
1973) are probably introgressive hybrids, with variable
proportions of wildcat genes (Kitchener and Easterbee
1992). Black forms (melanistic) have never been recorded
in wildcats in Europe, despite being a common coat color
mutation in other species of felid (Clark 1976, Robinson
1976, Todd 1977).
As with other wildcats, rodents are the staple of theirdiet across most of their range (Lindemann 1953, Novikov
1962, Nasilov 1972, Sladek 1973, Conde et al. 1972, Ragni
1978, Habijan and Dimitrijevic 1979, Hewson 1983, Stahl
1986, Riols 1988, Fernandes 1993, Ionescu 1993).
However, rabbits comprise the major prey where theyoccur, as in central Spain (Aymerich 1982), and an agricultural area in northeastern Scotland (Corbett 1979). Birds(both passerine and ground-dwelling)
are of secondary
importance (B. Ragni, P. Stahl in Zitt. 1992). The composition of the diet shows only minor seasonal variations: rabbits or rodents are the major year-round food items. No one

Action Planning
Projects 8 l-83.

Europeansilvestris,
Schreber,(Turkey);

wildcat, Fe/issilvestris group
1775

Other Names
Forest wildcat (English); chat forestier, chat sauvage, chatsilvestre (French); Wildkatze (German); gato mantes, gato
silvestre (Spanish); vairi katu, antarayin katu (Armenian);diwa kotka (Bulgarian); ghjattu volpe (Corsican); kodka
divoka (Czech); Wilde kat (Dutch); tkis cata (Georgian);vadmacska (Hungarian); gatto selvatico (Italian); zbik
(Polish);gato bravo (Portugese);pisica-salbatica(Romanian); dikaja koschka (Russian); macka diva

110

Pat? I: Species

species of rodent is preferred (Stahl 1986), but wildcats
sometimes prey selectively on rabbits. In northeastern
Scotland, for example, juvenile rabbits were taken in thespring birth season, and adults in autumn-winter, whenmyxomatosis was most virulent in that age class (Corbett1979). Wildcats will also scavenge food and cache theirkills, especially in winter (A. Kitchener in Zitl. 1993).
In western Scotland, Scott et al. (1993) found that wildcats were predominantly nocturnal, travelling over 10 km
per night to forage on open ground near the coast or aroundfarms and villages, and resting by day in thickets or youngforestry plantations. Daytime activity is usually correlatedwith absence of human disturbance (Stahl 1986, Genovesi
and Boitani 1993).
Wildcats can live in very wet, swampy areas (usually
among the last types of habitat to be modifed by humans).
N.K. Vereshchagin (in Heptner and Sludskii 1972)
describes how, when lowland forest is seasonally inundated in the Caucasus mountains, wildcats live in trees forweeks, feeding on rats taking refuge there.

.
cl

range

Protected
speciesarea
occursqx

whereProtected
speciesArea

area
probablywhere

4. Eurasia,

Europeanwildcat

Em-us: (C) 2-8 days, in presence of males (Conde andSchauenberg 1969).
Gestation: (C) 63-68 days (in Hemmer 1976).
Litter size: (C & W) mean 3.4 (W: n=106; C: n=92; Stahland Leger 1992); range l-8 (Green 1991).
Age at independence: (W)
Wijngaarden 1976, Tomkies(Muntyanu et al. 1993).

4-5 months (Smit and
199 1); up to 10 months

Age at sexual maturity: (C) females lo- 12 months; males
9-10 months (Conde and Schauenberg 1969, Hemmer
1976, P. Andrews in litt. 1993).
Interbirth interval: (C) probably one year. Conde andSchauenberg (1969, 1974) found that males can be sexually active from December-July, but females can only
exceptionally breed twice in one year, such as when thefirst litter is lost (A. Kitchener, B. Ragni in Zitt. 1993).
Mortality: (W) Several studies have reported very highhuman-caused mortality (e.g., snares, road kills), comprising up to 92% of observed deaths (Corbett 1979,
Piechocki 1986, Riols 1988). Human-induced mortalitv

whereoccursdistribution

Chapter

Wijngaarden 1976, Muntyanu et al. 1993).

Biology
Reproductive seuson: (W) mating season in late winter,
January-March;most births in May (Smit and Van

SpeciesAccounts.

IS uncertain

Figure 9. Distributionof the European
wildcat(F. silvestris,
silvesfris
group)
after Stahl andArtois (1991).
1. Montezinho
V (Portugal);
2. Nordeifel V; 3. Pfalzerwald
V (Germany);
4. Tatransky
II; 5. Beskydy
V (Poland);6. Hortobagyi
II* (Hungary);
7. Apuseni V (Romania);
8. Karpatskiy
II(Ukraine);
9. Karamanbayiri
VI; 10. Golardi Sulun VI (Turkey);
11. Kavkaz
I* (Russia);
12. Borzhom
I(Georgia);
13. Dilijan I (Armenia);
14. Zakatal
I (Azerbaijan);
15. Kodry I (Moldavia);
16. Coto Donana
II* complex (Spain).

111

Part I: Species

Accounts.

Chapter

4. Eurasia,

Europeanwildcatis probably significant across much of the wildcat’s
ily settled range (Stahl and Artois 199 1).

heav-unhesitating acceptance of survey results, finding a highdegree of error (39%) among experts (zoologists, natural history museum curators, hunters, veterinarians,
game wardens and professional naturalists) asked to distinguish between specimens of European wildcat anddomestic cat.
According to P. Stahl (in Zitt. 1992), changes and trends
in distribution are not well documented in most countries(Albania, Greece, Luxembourg, Poland, Portugal, Spain,
Switzerland, Turkey, former Yugoslavia). In two countries, the species became extinct in the first half of the 20th
century (Austria, Netherlands). In several west European
countries (Belgium, France, Germany, United Kingdom),
range expansion following World War II has been documented, although this expansion has now either halted, or
continues at a very low rate. In these countries and in Italy,
the range of the wildcat is generally considered stable,
although local declines have been found in parts of
Scotland (Easterbee et al. 1991). There seems to havebeen little change in wildcat populations in most east
European countries (Bulgaria, Hungary, Romania), except
in the Czech and Slovak Republics, where they havedeclined (P. Hell in Zitt. 1993).
A marked decrease in historical range has taken place inmost of the former Soviet Union (Bannikov and Sokolov
1984, Belousova 1993, Muntyanu et al. 1993, Puzachenko
1993a). Wildcat populations are now found in three major
areas: the Carpathian mountains of Ukraine (Bondar 1987,
Turyanin 1988); the Kodry region of Moldova (Montyanu
et al. 1993); and the Caucasus mountain region between
the Black and Caspian seas (Belousova 1993, Puzachenko
1993a). The broad-leaved forest habitat of the Ukrainian
Carpathians has been reduced by three- or four-fold overthe last century (Tatarinov 1983). These easternmost silvestris populations are important because the level ofhybridization with domestic cats is considered to be quite
low (Heptner and Sludskii 1972; see discussion underPrincipal Threats below).
In northeastern France, Artois (1985) found that wildcats used daily ranges of 0.3-3.3 km2. In the same study
area, Stahl et al. (1988) found that seasonal home ranges ofadult males were larger (5.7 * 2.6 km2; n=17) and morevariable in size than those of females (1.8 t 0.5 km’; n=7).
Resident male ranges overlapped 3-5 female ranges, butlittle overlap occurred between individuals of the samesex. In northeastern Scotland, however, Corbett (1979)
found that males and females had equivalent averagemonthly home ranges (1.75 km”), with little overlap. In
western Europe, densities of 3-5 animals per 10 km2 arereported from optimal forest habitats (review by Stahl andLeger 1992).
Stahl and Artois (199 1) reviewed the results of several
reintroduction attempts throughout Europe, and concluded

Longevity:- (C) up to 15 years (Green 1991).
Habitat and Distribution
European wildcats are primarily associated with forest,
and are found in highest numbers in broad-leaved or mixed
forests (Stahl and Leger 1992). Coniferous forest, however, is probably marginal habitat (Parent 1975, Heptnerand Sludskii 1972). Wildcats are also found in Mediterranean mayuis scrubland (Ragni 198 1), riparian forest,
marsh boundaries and along sea coasts (Lozan andKorcmar 1965, Heptner and Sludskii 1972, Dimitrijevicand Habijan 1977, Scott et al. 1993). They have neverbeen found in the high Alps (Schauenberg 1970); B. Ragni(in Zitt. 199 1) further states that forest wildcats are not present in areas where snow cover is greater than 50%, ismore than 20 cm deep, and remains for more than 100 daysof the year. In general, regions occupied by forest wildcats
are characterized by low human density, with cultivation
typically taking the form of grazing areas divided into
small plots. Rocky areas are a preferred micro-habitat(Heptner and Sludskii 1972). Wildcats are generally
absent from areas of intensive cultivation (Easterbee et al.
1991, P. Hell in Zitt. 1993).
After the marked decline of the forest wildcat and its
eradication from much of Europe between the late 1700s
and early 1900s recolonization has occurred since 19201940 in several countries (Belgium, Czech Republic andSlovakia, France, Germany, Switzerland, United Kingdom) (Stahl and Artois 1991 and in Zitt. 1991, 1993).
Populations of wildcats occur on Crete, Corsica, Sardinia,
and the Balearic Islands, as well as numerous other small
Mediterranean islands. Some authorities consider these
populations to be discrete subspecies, related most closely
to the Zybica group, and among the most endangered populations in Europe (Arrighi and Salotti 1988, Ragni 1988,
B. Ragni in litt. 1993). Vigne (1992), on the other hand,considers them to be feral forms of domestic cats introduced centuries before by humans.
Population Status
Global: Category 5c. Regional (Europe): Category 2.
IUCN: not listed. Stahl and Artois( 199 1) carried out acomprehensive status survey, using questionnaires andan extensive literature review, and Fig. 9 is based on theirwork. The authors have highlighted the importance of
establishing data collection networks, and praised theresults of such efforts in Scotland (Easterbee et al. 199 1)
and Hungary (Szemethy 1989). In Scotland, the method
appeared to be sufficiently sensitive to detect relatively
swift changes in the populations, as well as regional variation in status. However, Ragni (1993a) cautions against

112

Part I: Species

Accounts.

Chapter

4. Eurasia,

Europeanwildcatbelieved that less than 2% of the Belgian population can beconsidered hybrid animals. Randi and Ragni (1986, 199 1)
concluded, on the basis of electrophoretic analyses andmorphological data, that there is little probability ofgenetic flow between sympatric populations of forest wildcats and domestic cats.
Morphological and genetic studies of Scottish wildcats(Hubbard et al. 1992), on the other hand, point to frequenthybridization, although genetically distinct wildcats doremain in the remote areas of northern and western
Scotland. The Scottish wildcat (F.s. gmmpia) wasrecognized by Haltenorth (1957) asthe only valid subspeciesinthe silvestris group. Szemethy’s (1993) radiotelemetry
study in Hungary of sympatric wildcats (n=5) and feral
domestic cats (n=6) provides data on how hybridization
occurs and spreads. The feral cats’ home ranges were
smaller(0.8- 1.7km2) andlocatednearfarms; the wildcats’
home rangeswere larger (1.5-8.7 km2) and avoided thefarms. However, during the breeding season,male wildcats shifted their home rangesto cover the territories offemale farm cats. Szemethy (1993) also noted that someferal cats were able to live independently of the farms,
and adaptedto wildcat socialstructure.
Stahl and Artois (1991) recommendprioritizing investigations into the extent of hybridization in regionswhere
pastconditionswere conducive to it, i.e. where:that a long-term project run by the Bavarian Nature
Conservancy Association in Germany was the best.
Buttner (in press) states that 237 ( 136 males: 10 1 females)
captive-bred individuals were released from 1984 to 1993.
Although there has been evidence of population establishment and natural reproduction, released individuals suffered high mortality during their initial weeks in the wild
(due mainly to road kills), and the survival rate was estimated at about 30%. Stahl (1993) is of the opinion that,given the risks of hybridization, reintroduction should not
be considered a priority for wildcat conservation: efforts
should instead focus on protecting and supplementing
small isolated populations.
Protection Status
CITES Appendix II. National legislation: fully protected
over most of its range. Hunting prohibited: Armenia,
Austria, Belgium, Czech Republic, France, Germany,
Greece, Hungary, Italy, Luxembourg, Moldavia, Poland,
Portugal, Spain, Switzerland, Turkey, United Kingdom,
Ukraine.
Hunting regulated: Azerbaijan, Romania,
Slovakia. No legal protection (outside reserve areas):
Bulgaria, Georgia, Romania. No information: Albania,
Croatia, Slovenia, Yugoslavia (Stahl and Artois 199 1,
Ionescu 1993, Puzachenko 1993a, A. Bukhnicashvili in
Zitt. 1993). Hunting is permitted in Slovakia from 1 Dec28 Feb without restrictions.
Total harvest has declined
from about 900 in 1968- 1970 to about 160 in 199 1, and
P. Hell (in Zitt. 1993) recommends at least a five-year
moratorium to allow populations to recover.

lllPrincipal Threats
Hybridization between wildcats and domestic cats was
first reported almost 200 years ago (Bewick 1807), andhybrids have been observed throughout Europe (Stahl andArtois 199 1). However, the significance of the phenomenon is debatable. The importance of hybridization isdiminished if F. silvestris is considered a polytypic
species, and increased if the domestic cat is viewed as aseparate species. Significant progress is being made inEurope towards defining the felid “units of conservation,”
combining studies of morphology (including pelage characters) and genetics to clarify the relationship between
wildcats and domestic cats (Balharry and Daniels 1993,
Crovella et al. 1993, Fernandes 1993, Kitchener et al.
1993, Ragni 1993a,b, Puzachenko 1993b). It is likely thathybridization in Europe is more advanced than in otherregions of the wildcat’s wide geographic range.
Suminski (1962, 1977) believed that “pure” forest wildcats were essentially extinct in Europe, having comparedbiological and morphological criteria among a large number of specimens.
His findings have been disputed(Heptner and Sludskii 1972): Parent (1974), for example,llwild populationshave showna sharpdrop in numbers
over the pastfew decades;
wildcat colonization is recent;wildcat populationsare small andisolated;
humanpopulationdensity is markedly increased,with aconcomitant growth in numbersof domesticcats; andhabitat transformation(intensive agriculture and forest
plantations)is advanced.

Other threatshighlightedby the survey of Stahl andArtois(199 1) include habitat and population fragmentation; significant human-causedmortality, especiallyroadkills; andvulnerability to diseases
transmittedby feral cats. Clinicalexaminationsby McOrist et al. (1991) suggestthe possibility that feline leukemia virus (FeLV) occurs as a sustained infection in someScottish populations,rather thanas an occasional infection acquired from domestic cats.
FeLV is transmittedreadily amongyoung catsvia infectedbody fluids, such as during fighting or mating, and isalmost always fatal. B. Ragni (in Zitt. 1993)believes thatdiseasetransmission,rather than hybridization, is the more
seriousthreat resulting from the wildcat’s contact with its
domesticatedrelative.
Action Planning
Projects 10, 15, and 89-90.

113

Part I: Species

Accounts.

Chapter

5. The Americas

Part I
Species Accounts

Chapter 5
The Americas

Box 1
Vulnerability

Index to Species of the Region (in order of vulnerability)

Species

Habitat Association
St [Mar] (Tot) Score

Geog.
Range
(106 km2)

Score

Kodkod, 0. guigna
Andean mtn. cat, 0. jacobitus
Jaguar, P. onca
Oncilla, L. tigrinus
Margay, L. wiedi
Canada lynx, L. canadensis
Geoff roy’s cat, 0. geoffroyi
Puma, P. concolor
Ocelot, L. parcialis
Bobcat, L. rufus
Pampas cat, 0. colocolo
Jaguarundi, H. yaguarondi

N: 2 [2] (4)N: 2 [0] (2)
1:4[3](7)
N: 3 [I] (4)N: 2 [3] (5)
1:4[4](8)
I: 6 [I] (7)B: 8 [7] (15)
1:5[4](9)
B: 7 [4] (11)
B: 4 [6] (10)
B: 6 [4] (10)

R: 0.16
R: 0.62
M: 8.91
s: 2.90
M: 6.06
M: 5.06
S: 2.80
W: 17.12
W: 12.45
M: 7.24
S: 3.86
w: 13.53

-2
-2
0
-1
0
0
-1
+I
+I
0
-1
+I

-1

-1
0
-1
-1
0
0
+I
0
+I
+I
+I

Body
Size
Score
s
s
L
s
s
M
s
L
M
M
s
s

+I
+I
-1
+I
+I
0
+I
-1
0
0
+I
+I

Total
Score

Ranking

-2
-2
-1
-1
0
0
0
+I
+I
+I
+I
+3

1
1
2(A)
2
3
3
3
40
4
4
4
5

Key:
Habitat Association
St = number of strong + significant habitats
N = Narrow (-1); I = Intermediate (0); B = Broad (+I)
[Mar] = number of marginal habitats(Tot) = total number of habitats
Geographic
Range (in millions of km*)
R = Restricted (-2); S = Small (-1); M = Medium
Body size
L = Large (-1); M = Medium(0); W = Wide (+I)(0); Ss= Small (+1).(A) = Actively threatened

Regional Criteria:
Habitat association: Narrow = 2-5 habitat types; Intermediate = 7-9 habitat types; Broad = 1O-l 5 habitat types.
Geographic range: Restricted = _<I million km*; Small = 2-4 million km*; Medium = 5-9 million km*;
Wide = 14-17 million km?
Body size: Large = 35-135 kg; Medium = 7-20 kg; Small = 16.5 kg.

See the Introduction

to the Species

Accounts

for explanationof the vulnerability

114

rankingsystem(pp. 2-6).

Pat? I: Species

Kod kod, Oncifelis guigna(Molina, 1782)

Accounts.

Chapter

5. The Americas,

KodkodBiology
Gestation: (C) 72-78 days (P. Quillen in litt. 1993).
Litter size: (C) 1-3 (P. Quillen in litt. 1993); 3-4 (Housse1953).

Other Names
Guigna, chat du Chili (French); Chilenische Waldkatze,
Nachtkatze (German); gtiifia (Spanish); gato de Santa
Cruz, gato guifia (Argentina); huina (Chile).

Longevity: (C) up to 11 years (Weigel 1975).
Habitat and Distribution
Found only in Argentina and Chile (Fig. 1), the kodkod isstrongly associated with the moist temperate mixed forestsof the southern Andean and Coastal ranges, particularly
the Valdivian forest of Chile, which is characterized by thepresence of bamboo in the understory (IUCN 1992a). It
ranges up to the treeline at approximately 1,900 (Miller
and Rottmann 1976) to 2,500 m (Melquist 1984). In
Argentina, the kodkod has been recorded from moist montane forest which has Valdivian characteristics, including a
multi-layered structure with bamboo, and numerous lianas
and epiphytes (Dimitri 1972, N.O. Herr-era in Zitt. 1992).
Most records (nine out of 10 in Redford and Eisenberg
1992) coincide with the original distribution of temperatemoist Araucarian and Valdivian forest (37-48” S: Udvardy
1975, C. Weber in Zitt. 1993). J. Rottmann (in Zitt. 1993)
describes the kodkod’s habitat associations in order ofimportance as ( 1) evergreen temperate rain forest, (2)deciduous temperate moist forest, (3) sclerophyllous scrub,and (4) coniferous forest.
Sclerophyllous scrub occurs in central Chile, fromabout 30-37” S (Udvardy 1975). Only one specimen has
been collected from this habitat type, which is structurally
and faunistically quite different from the Valdivian forest.
That specimen (from the Valparaiso area, 33” S) was
described as paler than the Valdivian kodkods, with alarger skull and heavier dentition. Osgood (1943), whoclassified the specimen as a separate subspecies (0.~.
rnolinae), remarked on its similarity to the “salt desert”
race of Geoffroy’s cat., found in the Andes of northwestern Argentina, and speculated that further specimens mightarise to link the two cats, although this has not occurred. In
Chile, Geoffroy’s cats are known to occur only in theNothofagus beech forests of the far south, and this race ofGeoffroy’s cats bears little resemblance to the Valdivian
kodkod (Cabrera 196 1). It is puzzling that the kodkod
should show such marked differences in habitat association, implying a broad habitat selectivity, and still havesuch a restricted distribution, implying narrow selectivity.
Moreover, the kodkod is tolerant of altered habitats, being
found in secondary forest and shrub as well as primary forest, and on the fringes of settled and cultivated areas. C.
Weber (in Zitt. 1993) notes that the Valparaiso area, where
the kodkod is still present, has been settled for over 1,000
years, and was deforested, except for small remnant
patches, at least 150 years ago.

Description
and Behavior (Plate 6)
The kodkod is the smallest felid in the Americas, weighing
an average of 2.2 kg (n=3: Greer 1965a). It is a buffy tobrownish cat heavily patterned with small black spots.
The kodkod is closely related to Geoffroy’s cat, of which it
may be a subspecies- they do not appear to be sympatric(Hemmer 1978aj. The subject deserves further investigation, particularly in Argentina, where Geoffroy’s cat ismore common and widely distributed, and has been collected near to the kodkod’s known range (Redford andEisenberg 1992, O.N. Herrera in Zitt. 1992). In comparison
to Geoffroy’s cat, the kodkod has a small face and muchthicker tail (P. Quillen in Zitt. 1993). There is a high incidence of melanism (Cabrera and Yepes 1960, Osgood
1943, Greer 1965a) which, according to Miller andRottmann (1976), increases with latitude, and is particularly common on Chiloe and Guaitecas islands. The kodkod has rather large feet, and well-developed arboreal
abilities, sheltering in trees during inactive periods andclimbing as an escape tactic when pursued (Housse 1953,
Greer 1965a). Housse (1953) also notes that it dens inbamboo thickets.
Pearson (1983) examined the small mammal fauna of
the southern Andean moist temperate forest, focusing on
Argentina’s Nahuel Huapi National Park, where the kodkod is known to occur. He found that this forest type, withwhich the kodkod is strongly associated (see below), hasa high diversity of mouse-sized rodents, but lacks diversityof larger mammals, noticeably of the arboreal type (nosquirrels or monkeys). Most of the small rodents are terrestrial, semi-fossorial, and diurnal. Kodkod stomachs
have been found to contain remains of small rodents,
Norwegian rat, and birds (Koslowsky 1904, Housse 1953,
Greer 1965a). Kodkods have also been reported to takepoultry (Guggisberg
1975, Melquist 1984). While
believed to be primarily nocturnal (Miller and Rottmann1976), Green (1991) notes that most activity takes place
during the day in captivity.
The origin of the kodkod’s name is obscure. It may befrom one of the Mapuche Indian dialects, and probably
originally referred to the pampas cat (0. c&c&)-“colocola” may be a Spanish corruption of “kodkod” (F. Jaksicin Zirt. 1993). 0. gciigpza is most commonly called thegiiina (pronounced gwee-nya) in Chile and Argentina.

115

Part I: Species

Accounts.

Chapter

5. The Americas,

Andeanmountaincat1993). In the mid- 1800s the German naturalist Philippi
describedgroupsof kodkodsraiding chicken houses,with
up to 20 cats being killed by farmers in a single day(Cabrera and Yepes 1960, Guggisberg 1975). Farmers
there still complain of poultry depredationby the kodkod(Melquist 1984).

I

Protection Status
CITES Appendix II. National legislation:fully protectedin
Argentina andChile (Fuller et al. 1987).
Principal Threats
Becauseof its restricteddistribution, the kodkod is particularly vulnerable to habitat loss, the primary cause of
reducednumbersin the north of its range. However, there
is presently little forest clearancefor agriculturalpurposes,
and mostmonocultureplantationsarebeing establishedon
abandoned agricultural lands (C. Weber in ht. 1993).
Logging of the Chilean Valdivian forest is increasingfor
export to Japan(Ancient Forest International [AFI] 1990,
F. Jaksicin ht. 1993),but a substantialproportion (36%)
is protected(WCMC 1992:453), andlogging is not necessarily a threat to the kodkod becauseof its useof secondary
vegetation (J. Rottmann in ht. 1993). There are also several large protected areaswithin its range in Argentina(Melquist 1984,O.N. Herrerain ht. 1992). J. Rottmann(inZitt. 1993)mentionsfox hunting (both legalandillegal, withdogsandtraps)asapotential threat,citing unpublisheddata
showing that l-5% of fox hunter kills are small cats.
Melquist (1984), however, only once saw a garment ofwhat appearedto be kodkod pelts in a local market.

8.

Protected
area where
species
has been recorded

A

Confirmed

clclrecord3gure 1. Distributionof the kodkod(0. guigna).
1. Valparaiso
City (Osgood
1943, C. Weber in Mt. 1993);
2. Conguillio
II; 3. Puyehue
II; 4. Chiloe II; 5. Las Guaitecas
IV(Chile); 6. Lanin II complex;
7. Nahuel Huapi II complex; 8. Los
Alerces I I complex (Argentina).

Action Planning
Project 91.

Population Status
Global: Category 2. Regional: Category 1. IUCN:
Indeterminate. The kodkod has historically been described
as quite common (Osgood 1943, Cabrera and Yepes
1960). However, in the dry scrub of central Chile, 10%of the country’s area but home to two-thirds of its population (Weber 1983), habitat loss has led to localized andpatchy distribution (J. Rottmann in Zitt. 1993). In general,
however, the southern forested part of its range is well protected and sparsely populated by humans. Even where its
habitat has been altered, such as in central Chile, where
15,000 km2 of pine and eucalyptus plantations have been
established (C. Weber in Zitt. 1993), kodkods may do wellas rodent populations thrive (J. Rottmann, pers. comm. in
Melquist 1984).
The kodkod seems to live at higher densities on Chiloe
Island (Melquist 1984), possibly linked to the absence ofthe puma, grey fox, and Andean fox (J. Rottmann in Zitt.

Andean mountain cat,
Oreailurus jacobitus(Cornalia, 1865)
Other Names
Chat des Andes (French); Andenkatze, Bergkatze(German); chinchay, gato andino, gato lince (Spanish);gato mantesaltiplanico, titi (Bolivia); gato mantesandino(Chile); osjo (Peru).
Description and Behavior (Plate 6)
There arejust a few museumskinsand skullsby which todescribethis species,and there have beenjust a handfulof observations made in the wild (Grimwood 1969,
Scrocchi and Halloy 1986, Ziesler 1992). The Andean
mountaincat is a smallbut sturdy cat: onemalefrom Peru

116

Part I: Species

weighed 4 kg (Pearson 1957). It has long ash-grey fur
indistinctly patterned with rusty rosette-like spots on thesides, and marked with conspicuous thick dark stripes
extending down the sides from the back. Prominent darkgrey bars run also across its chest and forelegs. Its nose isblack, and its belly pale, with dark spots. The tail is thickand long (about 70% of head-body length: Osgood 1943,
Pearson 1957, Cabrera 1961, Pine et al. 1979), bandedwith approximately seven conspicuous dark rings. The
mountain cat is characterized by large auditory bullae of aform unique among the felids, the anterior chamber being
somewhat larger than the posterior (Kuhn 1973). Enlarged
auditory bullae are typical of animals inhabiting arid environments with little cover for protection and concealment(see description of the sand cat under North Africa andSouthwest Asia).
Essentially nothing is known about the biology andbehavior of the Andean mountain cat. The most detailed
observation of it in the scientific literature was made at
4,250 m in the northeast of Argentina’s Tucuman province(Scrocchi and Halloy 1986). A single cat was followed
on foot for more than two hours during late morning at a
distance of 15 to 50 m, showing no fear of humans. It
drank from melting ice, and moved to sit upon a prominentrock. A gray fox ran from the cat. The cat travelled further
and rested in the shadows on a rocky hillside before it
moved out of sight.
Halloy ( 1985) suggests that the Andean cat may be more
active on full moon nights, with more daylight activity during the new moon. Winters probably represent critical periods of increased hardship for the Andean mountain cat.
Burmeister (1879) reported without elaboration that the
Andean mountain cat prefers to hunt mountain chinchillas(nocturnal) and mountain viscachas (diurnal). Grimwood(1969) and Ziesler (1992) each observed a cat stalking
mountain viscachas at 4,000-4,300 m. These remain the
only clues to its diet, which may or may not include otherspecies (birds, reptiles, small rodents, etc.). The mountaincat’s range appears to coincide with the original distribution of these large rodent species. Both are “ricochettal”
rodents: their strategy to escape predators involves making unpredictable changes of direction by bounding off
rock faces (MacClintock 1966). Like the Andean mountain cat, the mountain chinchillas and viscachas haveenlarged auditory bullae. The long tail of the mountaincat (which is much longer than that of the similar-looking
montane form of the pampas cat [Redford and Eisenberg
19921) is probably an aid to balance when chasing theserodents. Other species with relatively long thick tails
include the cheetah (gazelles and hares change directions
swiftly during high-speed chase as an escape strategy), thesnow leopard (which hunts mountain goats and sheep
among cliffs and crags), and the clouded leopard, marbled
cat, and margay (species with highly developed arboreal
capabilities).

Accounts.

Chapter

5. The Americas,

Andeanmountaincat

Biology
No information. No animals known to be kept in captivity.
Habitat and Distribution
The Andean mountain cat is apparently very specialized
in its habitat requirements, having been found only in the
rocky arid and semi-arid zones of the high Andes above thetimberline (generally above 3,000-4,000 m in elevation).
Vegetation at observation and collection sites has consisted
mainly of small scattered dwarf shrubs and clumps ofbunchgrass (Pearson 1957, Scrocchi and Halloy 1986).
The presence of rock piles and boulders (typical microhabitat of mountain viscachas, and the only type of coveravailable at such altitudes) may be important (Scrocchi and

Altoandina
grassesandbiogeographicalherbs, 4,200m

Punefiabiogeographical(high grass plains andprovince(xerophyll
up to permanent

province
shrub steppes,

3,200

steppessnowline)of- 4,400m)

Figure 2. Distributionof the Andean
mountaincat (0. jxobitus).
Protected
areas where 0. jacobifus
may occur: 1. Lauca
II* complex (Chile); 2. Los Andes I; 3. San Guillermo
IV* complex(Argentina).
117

Part I: Species

Accounts.

Chapter

5. The Americas,

Jaguar

It is possible that the Andean mountain cat is rare
because it has evolved to be a specialized predator of chinchillids. Both mountain chinchillas and mountain viscachas have naturally patchy distributions, living in smallcolonies (the biggest viscacha colonies contain onlyaround 60 animals [Ziesler 19921). The colonies are centered around cliffs and boulders, and the animals avoid
extensive areas of open ground. Moreover, the highmountain habitat types are also unevenly distributed insome parts of the Andes, where the high plateau is dissected by deep valleys which are better watered, more
thickly vegetated and relatively heavily settled-not characteristic mountain cat habitat (Scrocchi and Halloy 1986).
More specifically, perhaps the Andean mountain cat
evolved to hunt nocturnal chinchillas rather than the larger,
diurnal viscachas (few cats are diurnal, and neither of thetwo observed hunts were successful). While mountain viscachas are declining locally outside of reserves due to subsistence hunting (H. Torres in Zitt. 1991, J. Rottmann in Zitt.
1993), the short-tailed chinchilla has been hunted to thebrink of extinction. It was intensively exploited for theEuropean fur trade from the late 19th to the early 20th centuries. Formerly ranging through the high Andes from
northern Peru south to the vicinity of Santiago, only a few
scattered colonies are believed to survive in rugged andinaccessible terrain where the borders of Argentina,
Boliva, Chile and Peru meet (at about 23” S) (Thornback
and Jenkins 1982, Gudynas 1989).
If the Andean mountain cat has indeed specialized toprey on chinchillas, widespread extinction of colonies
must have had disastrous effects. On the other hand, if it is
not a specialist predator, small prey biomass is high in the
Andean uplands (A. Canedi, C. Weber in Zitt. 1993), andits rarity must be attributed to the other factors.

Halloy 1986, Ziesler 1992). Figure 2 shows the area of
the Puna (grassland) and Altoandina high-altitude biogeographic provinces (Cabrera and Willink 1980), within therange delimited by 0. jcrcobitus collection and sighting
records (Grimwood 1969, Melquist 1984, Scrocchi andHalloy 1986, Redford and Eisenberg 1992). The distribution of these habitat types is more patchily distributed thanshown, as the high plateau is broken up by deep valleys.
Population Status
Global: Category 2. Regional: Category 1. IUCN:
Insufficiently Known. Andean mountain cats apparently
occur at low densities.
Protection Status
CITES Appendix I. National legislation: fully protected
over its range. Hunting and trade prohibited: Argentina,
Bolivia, Chile, Peru (Fuller et al. 1987).
Principal Threats
It is not clear whether the apparent rarity of the Andean
mountain cat is a natural phenomenon, is attributable tohuman actions, or is simply a misperception resulting fromlack of observations. Only a detailed study of its ecology
will provide the answer, but in the meantime, speculation
will have to suffice.
Lack of knowledge is obviously a factor. The few
observations of the species were all in the daytime, anddetails regarding collection or observation, typically madeduring general mammal surveys, are sparse (e.g., Pearson
1957, Greer 1965b, Grimwood 1969, Pine et al. 1979,
Melquist 1984). With regard to human action, it appears
that two of the usual human-induced causes of rarityhabitat loss/modification and direct persecution-are
onlypartially responsible.
There have been no significant
changes in land-use of the high Andes over the last 2,000
years-ifanything, the human population has decreased(S. Halloy in Zitt. 1993, C. Weber in Zitt. 1993). Grazing
by domestic camelids, sheep, and goats can lead to
reduced densities of large rodents, but at present this problem is localized rather than widespread (Holdridge 1978,
J. Rottmann in Zitt. 1993). Large dogs, feral or accompanying livestock, may chase and frighten Andean cats. In
terms of hunting pressure, herders usually carry guns
when accompanying grazing animals, and pelts of Andean
mountain cats are occasionally seen in fur markets(Melquist 1984, A. Ximenez in Zitt. 199 1, S. Halloy in Zitt
1993, J. von Thungen, pers. comm.). Yet C. Weber (inZitt. 1993) notes that the high Andes Indians of northern
Chile knew little of the species, and that all the pelts heobserved which were kept for ceremonial purposes wereof the pampas cat. There are no records of international
trade (aside from one probably misidentified transEuropean shipment in 1977: WCMC unpubl. data).

Action Planning
Project 92.

Jaguar, Panthera onca
(Linnaeus, 1758)
Other Names
Jaguar (French); Jaguar (German); tigre, tigre real, yaguar(Spanish); onca, onca pintada, onca cangucu (Brazil); tig
marque (French Guiana); yaguarete (Guarani); zac-bolay(Mayan); jaguarete (Paraguay); otorongo (Peru); penitigri(Suriname); yaguar (Venezuelan); onqa negra, yaguara
pichuna, yagua-hu (black jaguars).
Description
and Behavior (Plate 4)
The jaguar is the largest cat of the Americas, and the only118

Part I: Species

living representative of the genus Pnnthera found in the
New World. The jaguar’s pattern differs from that of theleopard by having larger, broken-edged rosettes aroundsmall black spots. It has a large head and stocky build,with relatively shorter limbs than others of its genus(Gonyea 1976). Melanism is frequent in the jaguar, andis inherited as a monogenic dominant to the normal
golden-colored form, rather than through a recessive allele(Deutsch 1975, Dittrich 1979). Albinistic specimens areoccasionally reported (Seymour 1989). Forest jaguars are
not only more frequently darker, but are also considerably
smaller in size than animals which inhabit more open
areas. In central American rain forest, 13 males averaged
57 kg and seven females 42 kg (Rabinowitz
andNottingham 1986, Aranda 1990), while in the Brazilian
Pantanal males averaged 100 kg (n=24) and females 76
kg (n=16) (de Almeida 1984). The size difference may
be due to the greater abundance of large prey species inmore open environments.
More than 85 species have been recorded in thejaguar’s diet (Seymour 1989). Large prey, such as peccaries, tapirs and deer, may be preferred, but a jaguar will
eat almost anything it can catch, and in the rain forest willtake mammal prey species in proportion to their occurrence (Rabinowitz and Nottingham 1986, Emmons 1987).
Large herbivores are more thinly distributed in rain forest
than in more grassy, open habitats, where they are more
likely to form groups and cluster near water, and jaguar
diet in the rain forest and in Savannah woodlands reflects
this difference in prey availability and vulnerability(Emmons 1991). In many areas, cattle are ranched onwhat is essentially prime jaguar habitat, and cattle havebeen the most frequent prey species documented in several
analyses of jaguar diet in Brazil (A. Almeida in
Hoogesteijn et al. 1993, Crawshaw and Quigley in prep.)and Venezuela (Hoogesteijn and Mondolfi 1992).
Jaguars are the only big cats which regularly kill prey(especially capybaras) by piercing the skull with theircanines (Schaller and Vasconcelos 1978, Mondolfi and
Hoogesteijn 1986, Crawshaw
and Quigley in prep.).
Emmons (1987) suggests that the massive head and stout
canines of the jaguar are an adaptation to “cracking open”
well-armored reptilian prey, such as land tortoises andriver turtles. She notes that, following the late Pleistocene
extinctions of large herbivores, the jaguar and the puma
were the only representatives of five genera of North
American felid to persist, and speculates that the jaguar
evolved to take advantage of a formerly super-abundant
prey base of water reptiles.
Although the jaguar has been characterized as primarily
nocturnal (e.g., Nowak and Paradiso 1983), radiotelemetry
has shown that they are often active during the daytime,with activity peaks around dawn and dusk. Jaguars havebeen found to be active for 50-60% of each 24-hour period

Accounts.

Chapter

5. The Americas,

Jaguar(Schaller and Crawshaw 1980, Rabinowitz and Nottingham 1986, Crawshaw and Quigley 1991). Crawshaw andQuigley (199 1) found that mean daily travel distance wassignificantly larger for a male (3.3+ 1.8 km) than forfemales (1.8 t 2.5 km). Both sexes tended to travel further
each day during the dry season. Rabinowitz and Nottingham (1986) found that radio-collared male jaguars tended
to remain within small areas (average 2.5 km’) for a weekat a time before shifting in a single night to other parts of
their range.
Biology
Reproductive season: (W) probably year-round, but
Rabinowitz and Nottingham (1986) reported that young
are usually born in the rainy season when prey is moreabundant, hence seasonal birth peaks reported in otherareas (e.g., January-April in Venezuela [Hoogesteijn andMondolfi 19921) may be correlated with prey availability.
Estrus: (C) 6-17 days.
Estrus cycle: (C) average 37 days, range 22-65 days
(Sadleir 1966, Stehlik 1971, Lea1 1979).
Gestation: (C) average 101, range 91 - 111 days (Hemmer
1976).
Litter size: (C & W) 1-4, mode 2 (Hoogesteijn
Mondolfi 1992).

and

Age at independence: (W) 1.5-2 years.
Age at sexual maturity: (C) 2-3 years, females; 3-4 years,
males (Mondolfi and Hoogesteijn 1986).
Longevity: (W) 1 l- 12 years (A. Rabinowitz, unpubl. data,
in Swank and Teer 1987); (C) up to over 20 years (Green
1991).
Habitat and Distribution
The jaguar, which swims well, is strongly associated withthe presence of water. Habitats meeting this requirement
range from rain forest to seasonally flooded swamp areas(Pantanal and Llanos), pampas grassland, thorn scrub
woodland (Chaco), and dry deciduous forest. In Belize,
Rabinowitz (199 1a) found that jaguars were more abundant in lowland areas of relatively dense forest cover withpermanent water sources than in open, seasonally dry
forests. In the Brazilian Pantanal, riparian forest wasstrongly preferred to open grassy areas (Crawshaw
andQuigley 199 1). Although jaguars have been reported fromelevations as high as 3,800 m (Costa Rica: Vaughan 1983),
jaguars typically avoid montane forest (Emmons 1991),
and have not been found in the high plateau of central
Mexico (Leopold 1959) or above 2,700 m in the Andes(Guggisberg 1975, Olrog and Lucero 198 1).
The historical range of the jaguar extended from119Part I: Species

Accounts.

Boundaries
in the early

Chapter

5. The Americas,

Jaguarof species
1900s

range

Populations

sustaining

or increasing

Populations

reducedor greatly

reducedlProtected

area where

species

occurs.

Protected
conserve

area over 3,000 Km2 in size which may
a minimum viable populationof 50+ jaguars

Current

Illl
El
cl
cl

Figure 3. Past and present
distribution
and relative
abundanceof the jaguar(/? once) after Swank andTeer (1987).
I. Calakmul
V* (Mexico)+
Maya IX* (Guatemala);
2. Montes Azules II (Mexico);
3. Cockscomb
Basin IV
(Belize); 4. Rio Platano IX#(Honduras);
5. La Amistad(Talamanca)
I I# complex(Costa Rica and Panama);
6. Darien II# complex(Panama)+ Los Katios II(Colombia);
7. Sierra Nevadade Santa Marta II* complex;
8. Sierra de la Macarena
IIcomplex;9. Cahuinari
II complex (Colombia);
IO. Yasuni II*(Ecuador);
11. Pacaya Samiria
VIII; 12. Manu II# (Peru);
13. Manuripi
Heath IV complex;
14. lsiboro Secure II (Bolivia);
15. Defensoresdel Chaco II(Paraguay);
16. lber IV complex (Argentina);
17. Mocone
Provincial
Reserve (Argentina)+ Turvo II (Brazil) complex;
18. lguazu II** (Argentina)+
Iguacu II** (Brazil) complex;
19. Jut-&a-ltatins
IV; 20. Alto
Ribeira II complex; 21. Pantanal Matogrossense
I I;
22. Araguaia
II complex;
23. Jau II complex (Brazil);
24. Pica da Neblina II (Brazil)+ Serrania de la Neblina II(Venezuela)
complex;25. Canaima
II; 26. AguaroGuariquito
II; 27. Henri Pittier
I (Venezuela).

range:

species, and densities may be as high as one resident per 15km”, as estimated for jaguars in Belize (Rabinowitz 199 1).
This refuge is of sufficient size and integrity to conserve
the species in large numbers for well into the forseeable
future, even if densities are lower than in Belize.
However, the jaguar is declining in most other habitats(Fig. 3). It has been virtually eliminated from much of
the drier northern parts of its range in the U.S. and Mexico(Brown 199 1), as well as the pampas scrub grasslands ofArgentina and throughout Uruguay (Swank and Teer
1987). The species has probably already lost significant
elements of its intraspecific diversity, and the trend continues. While commercial exploitation for their skins is nolonger a factor, jaguars still face local extirpation at thehands of cattle ranchers.

Arizona, New Mexico, and Texas in the United States
south to either the Rio Negro (40” S) or Rio Santa Cruz(50” S) in Argentina (Art-a 1974, Carman 1984). Formerly
occupied habitat in the north of its range included oak
woodland, mesquite thickets, and riparian forests (Brown
199 1). In the north, the jaguar’s range has receded southward about 1,000 km, and has been reduced in area by
about 67%. In South America, the jaguar’s range has
receded northward by well over 2,000 km, and has been
reduced by about 38% (Swank and Teer 1987).
Population Status
Global: Category 3(A). Regional: Category 2(A). IUCN:
not listed. The Amazon basin rain forest, some 6 million
km2 in extent (Collins 1990), is the key stronghold of the
120

--

Part I: Species

Accounts.

Chapter

5. The Americas,

Jaguarin significant numbers in an arid environment. Redford
et al. (1990) estimated that 176,000 km2 of viable habitat remain in the Paraguayan Chaco, an historical lossof 45%, with deforestation accelerating. Brooks (1990)
relays reports of Paraguayan biologists that jaguar populations are recovering from the depredations of theskin trade. Although the ecology of the species has notbeen studied, various density estimates for the region
have been put forward, ranging from one jaguar per 25
km2 to one per 75 km2 (in Swank and Teer 1987).

The status of several key large jaguar populations is discussed in detail below.
1. Yucatan peninsula/northern Guatemala/Belize: Rabinowitz and Nottingham(1986), working
in theCockscomb Basin of Belize, found male home ranges
to average 33 km2 (range 28-40 km2; n=4) and femalesa minimum of 10 km2 (n=3). There was partial overlap of male ranges, no overlap of female ranges, andmale ranges entirely encompassed those of females.
Based on a mean density of one resident adult jaguar
per 15 km’, and taking land and hunting pressures into
account, Rabinowitz (199 1a) estimated Belize’s population to number between 600 to 1,000. Based on density estimates (derived from footprints) of one jaguar
per 26-32 km2 in Mexico’s Calakmul Biosphere
Reserve, Aranda (1990) estimated a population of 125
180 jaguars for the 4,000 km2 reserve, and 465-550
jaguars in an adjoining 15,000 km2 of wilderness area
in Peten, northern Guatemala-which
has since beenprotected as the Maya Biosphere Reserve.

Protection Status
CITES Appendix I. National legislation: fully protected
over much of its range. Hunting prohibited: Argentina,
Brazil, Colombia, French Guiana, Honduras, Nicaragua,
Panama, Paraguay, Suriname, United States, Uruguay,
Venezuela. Hunting restricted to “problem animals”:
Brazil, Costa Rica, Guatemala, Mexico, Peru. Trophy
hunting permitted: Bolivia. No legal protection: Ecuador,
Guyana (Swank and Teer 1987, Fuller et al. 1987, Anon.
19894).

2. Chiapas state, Mexico: Aranda (in litt. 1993) estimated350 k 65 jaguars (based on a range of habitat-specificdensities from one animal per 15-40 km2) in the state,concentrated in four major populations occupying a
total of 8,800 km2.

Occurrencein Protected Areas:
Quigley and Crawshaw (1992) estimated that at least 3,200
km2 of protected habitat would be required to support aminimum population of 50 jaguars in the Pantanal region.
Protected areas over 3,000 km2 in size are marked with asquare in Fig. 3; other smaller but strategically locatedareas known to contain jaguar are also shown. Isolated
remnant populations are scattered through the fragmented
Atlantic coastal forests of southeastern Brazil, located inreserves (IUCN 1982) and also in unprotected areas,
including the Serra de Paranapiacaba Mountains southwestof Sao Paul0 (J. Guix in litt. 1993).

3. The Pantanal: The Pantanal is the largest seasonally
flooded land area in the world (Alho et al. 1988),
extending over 100,000 km2 on the borders of Brazil,
Bolivia and Paraguay. Wet Savannah woodland such as
this and the Venezuelan llanos may represen
represent t optimal
jaguar habitat, judging by the larger average sizes
attained by jaguars in these areas. However, the economy and culture revolves around cattle ranching in bothareas, and jaguar numbers have been greatly reduced.
Quigley and Crawshaw (1992) estimated jaguar density
at only 1.4 resident adults per 100 km2 in the Brazilian
Pantanal, where relatively undisturbed, intact populations exist only in the north-central and extreme southern parts of the region, separated by over 150 km.

Principal Threats
Deforestation rates are highest in Latin America (FAO
1993), and fragmentation of forest habitat isolates jaguar
populations so that they are more vulnerable to the predations of man. People compete with jaguars for prey
(Jorgenson and Redford 1993), and jaguars are frequently
shot on sight, despite protective legislation. The mosturgent conservation issue is the current intolerance ofranchers for jaguars (see Part II, Chapter 2 for more discussion of this issue). In many cattle-ranching operationsin the region, livestock roam widely and become essentially feral (Schaller 1983, Quigley and Crawshaw 1992).
Cattle have been shown to constitute a major portion ofjaguar diet in studies carried out on ranches in seasonally
flooded Savannah woodland (Hoogesteijn et al. 1993,
Crawshaw and Quigley in prep.). The vulnerability of thejaguar to persecution is demonstrated by its disappearance
by the mid- 1900s from the southwestern U.S. and northern
Mexico, areas which are today home to important puma

Resident jaguar annual home ranges averaged 142 km2(n= four females, one male)‘. The male’s home range
was not larger than the females’. Female home ranges
overlapped an average of 11.5% during the dry season,
but not the wet. Mean wet season home range (13 km’)
was significantly smaller than during the dry season
due to extensive flooding; jaguars used only 44-l13%
3% of
their total annual ranges during the wet (Crawshaw andQuigley 199 1).
4. Paraguayan Gran Chaco: The Chaco thorn scrub isprobably the only remaining place where jaguars occur

121Part I: Species

Accounts.

Chapter

5. The Americas,

Oncillalong (TL=26.9 cm, 56% of head-body length, n=13:
Redford and Eisenberg 1992). Melanistic individuals areoccasionally reported (Mondolfi 1986, Eisenberg 1990,
P. Quillen in Zitt. 1993). Prey taken from stomachs (n=3)
has consisted of small mammals (rodents and shrews) anda passerine bird (Gardner 197 1, Mondolfi 1986). Oncillas
have been reported to prey on small primates in Brazil (P.
Quillen in litt. 1993). J. Guix (in litt. 1993) analyzed thecontents of one stomach and five scats from Brazil’s
Atlantic coastal rain forest, and found feathers and hairfrom small mammals (rats and possibly one mouse opossum). Four oncillas were captured there in unbaited live
traps set along armadillo trails.

populations (Brown 199 1). A conservation plan has beendeveloped for jaguars in the Brazilian Pantanal (Quigley
and Crawshaw
1992), and the Brazilian government isplanning to establish a National Center for Research,
Management, and Conservation of Predators in Brazil toaddress livestock-predator problems (P. Crawshaw, pers.
comm.). Swank and Teer (1988) emphasize the potential
benefits of controlled sport hunting as an element of
national jaguar conservation strategies, arguing that trophy
fees would be an incentive for some ranchers to maintainjaguars on their land. Translocation of problem jaguars
has also been recommended (Anon. 1992c, 1993~).
Preliminary results from one such attempt in Brazil havebeen good (P. Crawshaw in litt. 1993), but Rabinowitz(1986) found that translocated jaguars in Belize often
returned to stock killing.
Commercial hunting and trapping of jaguars for theirpelts has declined drastically since the mid- 1970s whenanti-fur campaigns gathered steam and CITES controls
progressively shut down international markets (see Part II
Chapter 4). Organized poaching rings, in which fur buyers
travelled through the country supplying traps and buying
pelts from local people, are a thing of the past (Swank andTeer 1987).

Biology
Gestation: (C) 55-60 days (Widholzer et al. 1991 - 75-78
days (Leyhausen and Falkena 1966).
Estrus: (C) 3-9 days, with older cats having shorter cycles(Foreman 1988).
Litter size: (C) 1-3, one most common (Leyhausen andFalkena 1966, P. Quillen in litt. 1993).
Longevity: (C) average 11, but up to over 17 years (Prater
et al. 1988, P. Quillen in litt. 1993).

Action Planning
Projects 6 and 93-95.

Habitat and Distribution
The oncilla shows a strong preference for montane cloudforest (Mondolfi 1986), in that it is found at higher elevations than the ocelot and margay. For example, Melquist(1984) reports that it is restricted to elevations above 1,500
m in Colombia, and has been found at up to 4,500 m,
approximately snowline.
Rodriguez and Paz y Mifio(1989) also note that it has only been collected from theAndean highlands in Ecuador, a zone where the puma andpampas cat occur. Most specimens collected in Costa Rica
(5 of 6) have been taken in cloud forest (Gardner 197 1,
Vaughan in press). 111eastern Brazil, Koford (1973)
remarked on its presence in the subtropical forest highlands, and J. Guix (in Zitt. 1993) reports it from early secondary forest and abandoned eucalyptus plantation at 600
m elevation, in areas close to human settlement and highly
affected by deforestation and fire. Oliveira (1994) reported oncillas from semi-arid thorny scrub in northeast Brazil.
Bisbal(1989) also notes records from dry deciduous forest
in northern Venezuela. To what degree the oncilla uses
lowland moist forest is not clear, especially within the
Amazon basin. It has not been reported from this area (J.
Eisenberg in Zitt. 1993) and, according to L. Emmons (inZitt. 1993), is unlikely to occur there. However, there areseveral records from the outer edges of the Amazon rain
forest (Fig. 4), mainly from riverine forest (Mondolfi 1986,
Eisenberg 1990).
The oncilla appears to have a naturally disjunct distribution, although further research is necessary to confirm

Oncilla, Leopardus tigrinus(Schreber, 1775)
Other Names
Little tiger cat, little spotted cat (English); chat tigre,
oncille, (French); Onzille, Kleinflekenkatze, Ozelotkatze,
Zwergtigerkatze(German); tigrillo, tirica, gato tigre(Spanish); gato tigre Chico, gato onza Chico, gato pintado
Chico (Argentina); gato do mato (Brazil); chivi (Argentina,
Guyana, Peru); tigrillo peludo, tigre gallinero (Colombia);
caucel (Costa Rica); tigrillo Chico (Ecuador); chat tigre
tachete, chat tig (French Guiana); ocelot-cat, tigrikati(Suriname); tigrito (Venezuela).
Description
and Behavior (Plate 5)
With a silhouette and footprint resembling a house cat,the oncilla is small, having an average weight of 2.2 kg(n=3: Redford and Eisenberg 1992), with males slightly
larger than females (Guggisberg 1975). The oncillaclosely resembles the margay, and the two can be difficult
to distinguish in the field (Vaughan 1983). The two cats
are similarly marked, but the oncilla’s pattern of rosettes
tends to be less dark and blotchy than the margay’s, its fur
is not as thick, its body is more slender, and its tail not as122

Pat? I: Species

Accounts.

Chapter

5. The Americas,

Oncillathis. The northernmost record is from northern Costa Rica,
near the Tapand Cloud Forest Fauna1 Refuge (Vaughan inpress). It has been recorded from northern Panama(Melquist 1984), but the remainder of the country appears
to be a gap in the species’ range (Eisenberg 1990). Gardner(197 1) commented on the similarity of appearance of Costa
Rican cats to one collected in Colombia, and this moved
Melquist (1984) to state that the oncilla is probably foundthroughout Panama, as habitat there is suitable. There are
only two museum specimens for Ecuador and Peru
(Rodriguez and Paz y Mifio 1989, V. Pacheco in Z&t. to L.
Emmons). There are no museum records for Bolivia,
although K. Cassaro has reported seeing captive animals
originating from that country (pers. comm. to P. Quillen
1992). Figure 4 shows potential distribution if the oncillais assumed to be absent from the Amazon basin, and otherwise present only in montane and subtropical forest.
Population Status
Global: Category 3. Regional: Category 2. IUCN:
Insufficiently Known. This species has never been studied in the wild, and there is little understanding of its habitat requirements, density, and coexistence with othersmall cats. It has been trapped in large numbers for the furtrade-in
197 1, 28,000 pelts were counted in Brazilian
warehouses, and in 1983, 84,500 skins were exported
from Paraguay (Broad 1988)-although
it is likely that
other spotted cat pelts were mixed with oncilla. It is rarely
seen by field biologists.

la
cl
cl#
cl
.

A

Known
Protected
has been

and possiblearea whererecorded

speciesProtection
StatusUpgraded to CITES Appendix I in 1989. National legislation: protected over part of its range. Hunting prohibited: Argentina, Brazil, Colombia, Costa Rica, French
Guiana, Paraguay, Suriname, Venezuela. No legal protection: Ecuador, Guyana, Nicaragua, Panama, Peru
(Fuller et al. 1987).

range

speciesProtected
area where
species may occur
Confirmed

Principal Threats
It is difficult to assess threats to the oncilla when so little is
known about it. Coffee plantations are often established incloud forest habitat (Melquist 1984), but J. Guix’s (irz Zitt.
1993) observations of it in deforested areas and eucalyptus
monoculture on the outskirts of Sao Paolo suggest tolerance of habitat alteration. Although international tradeeffectively ceased after 1985 (WCMC unpubl. data), 675
spotted cat skins, mainly oncilla, were seized in Brazil, but
came from Paraguay. The age of the pelts, however, was
not ascertained (P. Crawshaw, A. Ximenez in Zitt. 1991).

recordFigure 4. Possible
distributionof the oncilla (L. tigrinus).
1. Tapanti IV (Costa Rica: Vaughan
in press); 2. La Amistad II*(Costa Rica and Panama:
Melquist 1984); 3. Sierra Nevada II
complex; 4. El Avila II; 5. Canaima
II (R. Hoogesteijn
in litt. 1993);
6. Specimen collected near lsla Chicharral,
Rio Negro, Amazonas (Mondolfi1986) (Venezuela);
7. Record from Rio Caqueta,
lowland gallery forest (Colombia:
Eisenberg
1990); 8. Specimen
collected from C. llalo, 2,800 m (Ecuador:
Rodriguez
and Paz y
Mine 1989); 9. Specimen
collected from La Florida, Region Alto
Yurinaqui, Junin, 800 m (Peru: V. Pacheco in lift. to L. Emmons);
IO. Specimen collected from northern
Mato Gross0 (Pine et al.
1970); 11. Four specimens
collected from secondary
forest scrubon the outskirts of Sao Paolo (J. Guix in litt. 1993) (Brazil).

Action Planning
Projects 96 and 97.

123

Part I: Species

Accounts.

Chapter

5. The Americas,

Margay, Leopardus(Schinz, 1821)

Margaywiedi

captivity; during the daytime, they rest in trees (Petersen
1977a, Konecny 1989).
Biology
Emus: (C) 4- 10 days.

Other Names
Margay (French); Langschwanzkatze(German); tigrillo,
gato tigre (Spanish); gato pintado (Argentina, Peru,
Venezuela); tigrillito (Belize); gato mantes, gato de month(Bolivia, Uruguay); gato maracaja mirim peludo (Brazil);
pichigueta (Cent. Am.); caucel (Costa Rica, Honduras);burricon (Ecuador); mbaracaya (Guatemala); kuichua(Guyana); chat tig. chat margay (French Guiana); chulul(Mayan); huamburushu (Peru, Venezuela); cunaguaro(Venezuela); tigrikati, boomkat [tree cat] (Suriname).

Estrus cycle: (C) 32-36 days (Petersen 1977b, Paintiff andAnderson 1980, Mellen 1989).
Gestation: (C) 76-84 days (Paintiff and Anderson
Mellen 1989, P. Quillen in Zitt. 1993).

1980,

Litter size: (C) one (n=l7: Mellen 1989), sometimes two(Mondolfi 1986).
Age at sexucrll
sexual maturity: (C) first estrus for females at 6-10
months (Petersen and Petersen 1978).

Description
and Behavior (Plate 5)
The margay is easily confused with the ocelot and theoncilla. It is difficult to distinguish by its coat, which ispatterned with black-ringed rosettes and elongate blotches
on a tawny background. Its fur is thick and plush. The
margay is between the two in size, weighing an averageof 3.2 kg (n=4: Redford and Eisenberg 1992). In motion,
however, the margay displays exceptional climbing abilities. It has the ability to rotate its hind foot through 180”
(Leyhausen 1963). Thus equipped, it can run straight
down trees head-first like a squirrel, and may hang from abranch by one hind foot. In the early 1800s Maximilian
Prince zu Wied observed the margay climbing about on
dangling lianas in the forest (Weigel 1975). Petersen( 1977a) describes the acrobatics of captive margays on arope strung horizontally in their enclosure. The cats would
jump from a distance of 2 m with front legs and claws
extended, hit the rope at their belly region, somersault overto hang by the hind feet, and drop back to the ground. The
margay’s tail is proportionately quite long (although not aslong as the Andean mountain cat), averaging 70% of headbody length (TL=36.4 cm; n=6: Redford and Eisenberg
1992). It serves as a counterweight
to aid balance(Eisenberg 1990).
In Belize, a radio-collared margay was found to forage
in trees, but travel on the ground (Konecny 1989). Basedon analysis of 27 scats, the most common item in the diet
was a small arboreal mammal, the big-eared climbing rat.
Squirrels, opossums, arthropods, small birds, and fruit
were also taken (Konecny 1989). Other reported arboreal
prey includes prehensile-tailed porcupines, marmosets,
capuchin monkeys, and three-toed sloths (Goldman 1920,
Beebe 1925). Terrestrial prey has also been reported,
including spiny pocket rats, cane rats, and cavies(Carvalho 1958, Mondolfi 1986), but in the Belize study
margays took more birds and fruit and fewer terrestrial
mammals than ocelots or jaguarundis (Konecny 1989).
Margays are strongly nocturnal, with highest levels ofactivity recorded between 0100-0500 both in the wild andLongevity: (C) up to 20 years (Prator et al. 1988).
Habitat and Distribution
The margay is more strongly associated with forest habitat, both evergreen and deciduous (Bisbal 1989), than any
other tropical American cat (Fig. 5). Although it formerly
occurred as far north as Texas, it was restricted to riverine forest (L. Emmons in Zitt. 1993). It hunts on theboundaries of such gallery forests and may thus be foundsome distance from forest cover in Savannah habitats(Eisenberg 1990). It appears to be less tolerant of humansettlement and altered habitat than its close relatives, theocelot and oncilla, although it has been occasionallyreported outside forested areas (Vaughan 1983, Tello
1986b), such as in shady cocoa or coffee plantations in
Venezuela (Mondolfi 1986). In a successional forest
mosaic in Belize, one sub-adult male margay spent significantlv
nificantly more time in late secondary growth forest thanin other habitat patches, but would make 2-3 day forays
into early secondary growgrowthth forest (Konecny 1989). Johns(1986) reports, howhowever,
ever, that the margay is absent fromlogged areas. The margay is rarely found at elevations
above 1,200 m (Eisenberg 1990).
Population Status
Global: Category 4. Regional: Category 3. IUCN:
Insufficiently Known. Very little is known of the margay’s status and abundance across its range. The Amazon
Basin is its stronghold. It is reported to be rare in northwestern Argentina (Mares et al. 198 1, A. Canedi in litt.
1993) and Uruguay (A. Ximenez in Zitt. 1990). A subadult male in successional forest in Belize’s Cockscomb
Basin Wildlife Sanctuary-wheremargays appear to beless common than in primary forest-had
a home rangeof 11 km* (Konecny 1989). An adult radio-collared malein Brazil’s Iguacu National Park monitored over 18
months by P. Crawshaw (pers. comm. 1993) maintained ahome range of 16 km*.

124

Part I: Species

Accounts.

Chapter

5. The Americas,

Margayappear in international trade at a time of concern over thelevel of exploitation of the ocelot, and species of spottedcats in trade were rarely verified. Illegal hunting for domestic markets or for the underground skin trade has been reported to be a continuing a problem in some areas (Melyuist
1984, Paz y Mine 1988, Walton 1991). Aranda (1991)
reported that the margay’s pelt was the most common in theskin trade in the southern Mexican state of Chiapas, despite
its protected status. A margay skin sold for U.S. $5 10,
while an ocelot pelt could fetch U.S. $50-90. However,
deforestation is now the primary threat to reduced populations now that international trade has virtuallv ceased.

Protection Status
Upgraded to CITES Appendix I in 1989. National legislation: fully protected over most of its range. Hunting prohibited: Argentina, Brazil, Bolivia, Brazil, Colombia,
Costa Rica, French Guiana, Guatemala, Honduras,
Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname,Uruguay, Venezuela. No legal protection: Ecuador,
Guyana, El Salvador (Fuller et a2. 1987).
Principal Threats
The margay has been one of the most heavily exploited
Latin American cats, with an average annual net tradereported to CITES of 13,934 skins between 1976 and 1984
(Broad 1987). Trade statistics probably do not reflect theactual number of margays killed, as margays began todAction Planning
Projects 96 and 97.

Figure 5. Distributionofthe margay(L. wiedi).
1. Montes Azules II*; 2. Calakmul V* (Mexico);
3. Tikal II**
(Guatemala);
4. Cockscomb
Basin IV (Belize); 5. Rio Platano
II* (Honduras);
6. Santa Rosa II
complex (Costa Rica); 7. La
Amistad II* (Costa Rica andPanama);8. Guatopo
II; 9. Perija
I I (Venezuela);
10. El Cocuy II
complex;
11. Amacayacu
II(Colombia);
12. Cerros de
AmotapeII complex;
13. Pacaya
Samiria IV; 14. Manu II# (Peru);
15. Manuripi
Heath IV complex;
16. Noel Kempff Mercado
II(Bolivia); 17. Baritu II (Argentina); 18. Quebradade Los
Cuervos
reserve (proposed);
19. Santa Teresa V (Uruguay);
20. Iguacu II** (Brazil) + lguazu
I I** (Argentina)
complex;21. Caparao
II; 22. Amazonia(Tapajos)
II complex (Brazil).

Protected
has beenarea whererecorded

species125Part I: Species

Accounts.

Chapter

5. The Americas,

Canadalynx

Canada lynx, Lynx
canadensis Kerr, 1792
Other Names
Lynx du Canada (French); Kanadaluchs (German); lince
de1 Canada (Spanish).
Description
and Behavior (Plate 6)
The Canada lynx has a flared facial ruff, black ear tufts,
and long hind legs which lend a slightly stooped posture.
The pelage is reddish-brown to grey; the hairs are tippedwith white which gives the fur a frosted appearance. Thereis a rare pallid color phase which suggests partial albinism,
known as the blue lynx in the fur trade (Quinn and Parker
1987). The Canada lynx’s large spreading feet act likesnowshoes, and are twice as effective at supporting its
weight on snow as those of the bobcat (Parker et al. 1983).
The lynxes show remarkable similarity of appearancecompared to other related groups of cats, and the Canada
lynx is often treated as conspecific with the Eurasian lynx
(Kurten and Rausch 1959, Tumlison 1987). However, theCanada lynx is only half the size of the Eurasian lynx:
average adult weight of Canada lynx males is 10.7 kg(n=201) and females 8.9 kg (n=183) (U. Breitenmoser and
C. Breitenmoser-Wursten
in prep.). While the Canada
lynx is probably a descendent of a Eurasian lynx ancestor
which migrated into North America during one of the last
two major glacial periods (Werdelin 1981, 1983b), theBreitenmosers (in prep.) argue convincingly that the twoshould be considered separate species, as they now show
marked adaptive differences for prey capture. Whereas thelarger Eurasian lynx preys mainly on ungulates, theCanada lynx relies almost exclusively on snowshoe hares,
and is uniquely adapted, both behaviorally and physiologically, to exploit a cyclic prey base.
Among felid predator-prey relationships, there are none
as closely tied as that between the hare and the Canada
lynx (Van Zyll de Jong 1966, Nellis et al. 1972, Brand
and Keith 1979, Parker et al. 1983, Ward and Krebs 1985).
The lynx-hare cycle was first discovered from harvest
records of the Hudson’s Bay Company dating back to the1800s (Elton and Nicholson 1942: Fig. 6). Numbers ofsnowshoe hares peak approximately every ten years, andlynx numbers follow the same pattern with a short lag, typically one to two years (Keith 1963, Bulmer 1974). While
the populations of many prey and predator species are
cyclic and roughly synchronous in the northern latitudes,
the snowshoe hare and lynx correlation is particularly
close (Keith 1963, Mallory 1987: Fig. 7). The amplitudeof the lynx population cycle is greater than that of any
other predator (Bulmer 1974), and lynx density during
cyclic highs and lows can differ by up to 15fold(Breitenmoser et al. 1993b). As hares decline, fewer lynx

’
1823

1830

1840

1850

1860

*

L

I070

1880

*.
1890

*.
1900

I910

1920

1930

19-a

YEAR

Figure 6. The IO-year
cycle in Canada
lynx populations
visible in annual recordsof lynx pelt purchasesby the Hudson
Bay Company,
Canada.
Source: van Zyll de Jong (1971).

breed, producing smaller litters with few, if any, surviving kits. As hares increase, so do lynx reproduction andrecruitment rates (Nellis et al. 1972, Brand and Keith
1979, Parker et al. 1983, O’Connor 1984, Slough andWard 1990, Breitenmoser et al. 1993b, Mowat 1993). In
captivity, female lynx do not show such an early onset of
sexual maturity or such high litter sizes as during hare
peaks in the wild (Breitenmoser et al. 1993b). While lynx
will switch prey during periods when hares are scarce(Brand et al. 1976), turning to small rodents, ground birds
and, exceptionally, ungulates such as white-tailed deer,
caribou, and Dal1 sheep (Saunders 1963, Bergerud 1983,
Stephenson et al. 1991), lynx populations only reach highdensities when supported by snowshoe hares (Brand andKeith 1979, Mech 1980, Ward and Krebs 1985).
There are several competing hypotheses to explain thehare cycle. The most widely accepted explanation is thatwinter food shortage (Keith 1974) depresses hare reproduction (Carey and Keith 1979) at the population peak andstarts the cyclic downturn, and hare numbers are subsequently further reduced due to predation (Keith et al. 1984,

126

Pati I: Species

Interbirth interval:(Tumlison 1987).

Mortality rates: (W) adult mortality rates average 55%
for exploited populations in Canada (R. Ward in prep.
cited in Slough and Ward 1990). Rates vary dramaticallywith the hare cycle. Poole (1994) estimated survival rates
in an unharvested population to be 90% before and during
the decline in hare densities; 25% during the first year of
low hare densities; and 37% during the second year of thelow. All radio-collared lynx resident prior to or during
the hare decline dispersed and/or died by the end of thefirst winter of low hare densities.
Longevity: (W) up to 15 years (Nava 1970; K. Poole, B.
Slough unpubl. data).
Habitat and Distribution
Lynx are distributed throughout the broad boreal forest
belt of North America (Banfield 1974) and south into theAmerican Rocky Mountains (Koehler 1990b), with a total
range of some 7.7 million km2 (Parker and Quinn 1987:
Fig. 8). The historic range is largely intact, although it
has shrunk in the south due to human settlement and forest clearance (Banfield 1974, Quinn and Parker 1987).
Lynx will inhabit farming country, but only if it is interrupted by sufficient areas of woodland (Todd 1985).
Bobcats appear to be expanding northwards, and have displaced lynx in some areas (Parker and Smith 1983, Rolley1987). Lynx will travel long distances during both phasesof the hare cycle seeking out patches of hare abundance(Ward and Krebs 1985), with movements of up to 1,200
km recorded (K. Poole, B. Slough & G. Mowat, unpubl.

-

1 1

LYNX

l

\

X-O

\

l, l .

.\
‘0

CT3=.

.

l

.--a
.,.---

‘m

\

LL

OY

,

T

I

,

1932

36

40

44

/
w

9

48

T

I

1

52

56

60

(W) generally one year, rarely two

Recruitment rates: (W) 60-80% when hares are abundantand increasing, and approaching zero during lows (Brand
et al. 1976, Brand and Keith 1979, Parker et aI. 1983,
Mowat 1993, Poole 1994). Koehler (1990a) found low
recruitment rates of around 12% from 1980- 1987 inmature forest in north-central Washington (sub-optimal
habitat at the southern edge of lynx range).

Litter size: (W) higher (average 3.8-5.3) when prey isabundant, and reduced (2.3-3.5) when prey is scarce(Brand and Keith 1979, Slough and Ward 1990, Mowat
1993); range l-8 (Tumlison 1987, Breitenmoser et al.
1993b). Yearling lynxes give birth to smaller litters (O-4.2:

L-

lynx

Reproductive rates: (W) up to 100% during hare peaks,
and as low as zero during cyclic lows (Mowat 1993).

Gestation: (W) 63 to 70 days (Saunders 1961).

\

CanadaAge at first reproduction: (W) 10 months (first winter)when prey is abundant, more generally 22-23 months (second winter) (females); second year (males) (Saunders
1961, Van Zyll de Jong 1963, Stewart 1973, Nava 1970,
Brand and Keith 1979, O’Connor 1984).

Estrus: no published information on duration or length ofcycle (G. Mowat et al. in Zitt. 1993). It is possible that
ovulation in the Canada lynx may be at least partly spontaneous, although this is controversial (Van Zyll de Jong
1963, Quinn and Parker 1987, G. Mowat et al. in litt.
1993). Lynx may be induced ovulators when prey density
is low and there is less chance of meeting a mate, and spontaneous ovulators when prey density is high, improving
prospects for breeding and raising young (Kitchener 199 1).

0-m

5. The Americas,

Age at independence: (W) generally around 10 months:
kits typically leave their mother’s range in March-April(Saunders 1961, Bailey et al. 1986, Slough and Ward
1990, Mowat 1993).

Biology
Birth season: (W) May-June, exceptionally July (Saunders
1961, Nava 1970, Nellis et al. 1972, Mowat 1993).

I

ChapterMowat 1993)

Boutin et al. 1986). Gilpin (1973) and Schaffer (1984)
modelled harvest data mathematically, and concluded thatthe cycle is more complex than a simple predator-preyinteraction, involving at least a third additional factor.
Another suggested influence involves changes in the nutritional quality of vegetation in response to hare browsing(Bryant 198 1, Sinclair and Smith 1984, Bryant et al.
1985). Nevertheless, in some areas, hares have declined
even when food resources appear sufficient (Keith et al.
1984, Krebs et al. 1986). Preliminary results achieved
from long-term field experiments (Krebs et al. 1992) now
favor the hypothesis that predation alone, by a variety of
specialist and generalist carnivores, is the driving force
behind the cycle, as has been suggested for microtine
rodents (Hanski et al. 1991).

U-J
z
CL3
E

Accounts.

YEAR

Figure 7. The relationshipbetweensnowshoehare andCanada lynx, showing
the lag between
the two population
cycles.
Source: Keith (1963), from Mallory (1987).

127

Part I: Species

Accounts.

Chapter

5. The Americas,

Canadalynx

Figure 8. Distributionof theCanadalynx (L. canadensis).
1. Arctic IV complex; 2. Noatak
II* complex;
3. lnnoko IV;
4. Lake Clark II; 5. Katmai II;
6. Kenai IV complex;
7. Denali
II*; 8. Wrangell-Saint
Elias II;
9. Glacier Bay II; IO. North
CascadesII complex;
11. Glacier II*; 12. Yellowstone II# complex (U.S.).

Protected area
species occurs
Protected area where
species probably occurs
Extralimital records (Banfield 1974,
Govt. of Canada 1988, Koehler 1990b)
Extirpateddata). Extralimital records have documented lynx in thenorthern tundra and Arctic islands (Banfield 1974, Govt.
of Canada 1988), and in Iowa, South Dakota, Nebraska,
and West Virgina (Koehler 1990b). Snowshoe hares prefer new growth vegetation, such as after forest fire or logging, and lynx may cluster in these areas (Quinn andParker 1987).

New York (reintroduced), Vermont, New Hampshire, andMaine, with the largest populations in the Rocky
Mountains. Washington State recently listed the lynx as
Threatened, and will take more active measures to aid population recovery (Anon. 1994b). Much of the lynx’s
American range consists of National Forest lands (Koehler
1990b).
Lynx density fluctuates dramatically with the hare
cycle (Breitenmoser et al. 1993b). An ongoing long-term
study of an unexploited population in good quality habitatin the Yukon found densities of 2.8 individuals (including
kittens) per 100 km2 during the hare low, and 37.2 per 100
km2 during the peak (G. Mowat and B. Slough, unpubl.
data). Poole (1994) obtained very similar figures for his
study area in the Northwest Terrritories: 30 lynx per 100
km2 at the peak, and around 3/100 km2 the winter following the hare crash. In the south of their range, where snowshoe hare populations appear to be non-cyclic and stable at
low densities, Koehler (1990a) reported lynx density at 2.6
individuals per 100 km2 (north-central Washington). The
study was conducted in mature coniferous forest where
fires had been suppressed, and the early successional
growth preferred by snowshoe hares was limited to isolated pockets.
Home range sizes for lynx range from 4-25 km2 forfemales, and 4-70 km2 for males (G. Mowat and B.
Slough, unpubl. data). On the Kenai peninsula, Alaska,

Population Status
Global: Category 4. Regional: Category 3. IUCN: notlisted. The status of the lynx is generally satisfactory(Quinn and Parker 1987, Govt. of Canada 1988). In
Canada, it is considered endangered only in New
Brunswick, and has been extirpated from Prince Edward
Island and mainland Nova Scotia. The largest populations are found in southern Quebec, northern Alberta,
northern British Columbia, Yukon, the Northwest
Territories and Alaska (Govt. of Canada 1988; K. Poole,
B. Slough in Zitt. 1993). There is some concern that trapping pressure during the 197Os- 1980s may have reduced
population levels (see Part II Chapter 4), but R. Eagan(pers. corm-n.) draws attention to low hare cycles since theearly 1970s.
The main U.S. lynx population is found in Alaska.
Elsewhere, they are more sparsely distributed, occurring in
low numbers in the states of Washington, Montana, Idaho,
Wyoming, Colorado, Minnesota, Wisconsin, Michigan,

128

Part I: Species

Accounts.

Chapter

5. The Americas,

Geoffroy’scatquota system as lynx numbers increase.
Government authorities have either implemented theserecommendations or initiated harvest impact research programs, as discussed in Part II, Chapter 4. In addition, trapping methods may change as a result of pressure from theEuropean Union for countries to ban the use of the leghold
trap and adopt internationally accepted humane trapping
standards (see the bobcat Species Account).
Quinn and Parker (1987) do not believe that habitat
alteration has had significant impact on lynx populations,although in the southern portions of its range optimal habitat for snowshoe hares is more patchily distributed (Wolff
1980, Sievert and Keith 1985). Modified logging, leaving interspersing areas of good tree cover, can actually
benefit both lynx and their prey (Koehler and Brittell
1990). However, suppression of forest fires limits early
successional growth favored by hares (Fox 1978), and may
ultimately reduce hare abundance (B. Slough in Zitt. 1993).

Kesterson (1988) found larger home ranges-l 07 km2 forfemales and 225 km2 for males-but seasonal ranges were
smaller, with females only 9.4 km2 in summer. Male
ranges usually encompass those of females (Saunders
1963, Berrie 1973, Parker et al. 1983, Ward and Krebs
1985, Kesterson 1988, Slough and Ward 1990), but samesex overlap has also been found (Berrie 1973, Mech 1980,
Carbyn and Patriquin 1983, Noiseux and Doucet 1987; G.
Mowat, B. Slough and K. Poole unpubl. data). Breitenmoser et al. (1993b) suggest that same-sex overlap reflects
a high degree of tolerance of independent offspring byresident lynx, another unusual adaptation of the Canada
lynx to a predictably cyclic prey base.
Protection Status
CITES Appendix II. National legislation: managed for
exploitation over most of its range. In Canada, trapping
is regulated through closed seasons, quotas, limited entry
and long-term trapping concessions. (See Part II Chapter
4 for a more detailed discussion of harvest management.)
In the United States, trapping is permitted only in Alaska,
Idaho, and Montana (Koehler 1990b).

Action Planning
Project 98.

Principal Threats
In general, the future of the lynx looks more promisingthan for many other felids. However, there is still cause forconcern, as harvests of Canada lynx during the cyclic low
periods have progressively fallen since the mid- 1970s andhare numbers have similarly been lower since these periods (Todd 1985, Govt. of Canada 1988, R. Eagan, pers.
comm.).
Lynx are easily trapped in comparison to other furbearers (Quinn and Parker 1987). At the low point of the hare
cycle, lynx may become more vulnerable to exploitation as
they disperse in search of food-travelling
greater distances can increase the chances of being caught in a trap.
Recruitment is also falling during this phase of the cycle,
and it is possible that trapping pressure could reduce numbers to the extent that recovery to previous levels is not
attained when hares again increase (Brand and Keith 1979,
Parker et al. 1983, Bailey et al. 1986).
Several management options have been recommended
to prevent over-trapping, including prohibiting exploitation in hare refugia (small patches of optimal habitat)
throughout the cycle (Slough and Ward 1990, Poole 1992).
In the past when lynx pelt prices were high (U.S. $685 in
198 1), trappers would seek out these refugia and concentrate their trapping efforts there (Carbyn and Patriquin1983). Brand and Keith (1979) recommended that harvests be completely suspended for the 3-4 year low of thehare cycle, so that potentially more lynx are available for
harvesting in peak years. Bailey et al. (1986) recommended a combination of harvest suspensions in the moreaccessible trapping areas during low hare years, and a

Geoff roy’s cat, Oncifelis
geoffroyi (d’orbigny andGervais, 1844)
Other Names
Chat de Geoffroy (French); Geoffroykatze,
Kleinfleckkatze, Salzkatze (German); gato de mato, gato montes, gato de las salinas (Spanish); gato mantes comun(Argentina); gato do mato pelo curto, gato do mato deGeoffroy (Brazil); gato mantes argentino (Chile).
Description
and Behavior (Plate 6)
Geoffroy’s cats are small cats, uniformly patterned withsmall black spots of nearly equal size and spacing. Coatcolor tends to ochraceous in the north of its range and greyin the south (Guggisberg 1975, Ximenez 1975). Melanism
is fairly common (Cabrera and Yepes 1960, Foreman
1988, Brooks 1992). Adult males in Patagonia weighed anaverage of 4.8 kg (n=5), and females 4.2 kg (n=2)(Johnson and Franklin 1991). Geoffroy’s cats have been
described as good swimmers that readily enter water(Weigel 1975). In southern Chile’s Torres de1 Paine
National Park, one female was known to have crossed a 30
meter wide fast-flowing river at least 20 times (Johnson
and Franklin 199 1). Fish (Characidae) were found in an
examination of stomach contents in Uruguay, in addition
to amphibians, reptiles, birds, and small mammals (A.
Ximenez in Zitt. 1990). Geoffroy’s cats are also described

129

Pat? I: Species

Accounts.

Chapter

5. The Americas,

Geoffroy’scat

1973, Melquist 1984, Brooks 1992), with the exception ofsouthern Chile, where it is restricted to a small area of cold
scrublands east of the Andes (Glade 1988). However,there are fears that a decade of high-volume skin trade has
severely reduced populations (Govt. of Brazil 1992). Its
status is not well known.
In Torres de1 Paine National Park, female annual homeranges averaged 3.7 t 1.1 km2 (n=2), and males 9.2 t 1.7
km2 (n=5) . Female ranges overlapped, while males apparently did not. Density was estimated at 1.2 individuals
per 10 km2 (including kittens) in prime habitat (Notho-

as primarily nocturnal and partially arboreal (Cabrera andYepes 1960). In the first radiotelemetry study of thespecies, Johnson and Franklin (199 1) confirmed thisdescription: Geoffroy’s cats rested during the daytime
either in dense ground vegetation or in tree cavities. Of
325 scats located by the researchers, 93% were in arboreal middens in the crooks of trees. The diet consisted
primarily of rodents and European hares. Hares wereselected especially during the spring and summer birth season when juveniles were abundant. Geoffroy’s cats weretwice seen carrying European hare carcasses into a tree.
Biology
Birth season: (W) December-May(Ximenez
Johnson and Franklin 199 1, Brooks 1992).

1975,

Estrus: (C) 2.5 t 0.5 days (n=2).
Estrus cvde:(C) 20 days (n=l: Mellen 1989).
i
Gestution: (C) 72-78 days (Green 1991).
Litter size: (C) 2.3 1 k 0.13 (n=14) (Mellen 1989).
Age at sexual maturity: (C) 18 months (females), 24
months (males) (Foreman 1988), but as early as 9- 12
months (P. Quillen in Zitt. 1993).
Interbirth intend:(W) reportedly(Cabrera and Yepes 1960).
Longevity:1993).

one litter per year(C) up to over 14 years (P. Quillen in Zitt.

Habitat and Distribution
Geoffroy’s cat has been described as occurring in a wide
variety of habitat types (Koford 1976, Melquist 1984,
Broad 1987). It is distributed throughout the pampas
grasslands and arid Chaco shrub and woodlands, and up
around the Salinas Grandes (alpine saline desert of northwestern Argentina) to 3,300 m in the Andes (Cabrera andYepes 1960, Ximenez 1975, Brooks 1992). However, it
is not found in either the tropical rain forests or southernbroad-leaved forests, and avoids open areas, preferring
dense, scrubby vegetation (Burmeister 1879, Johnson andFranklin 199 1). Therefore, throughout the broad stretchesof Southern Cone grassland, it, is restricted to forest
patches or riparian vegetation (Cabrera and Yepes 1960,
Ximenez 1975). It is sympatric throughout its range withthe pampas cat, and the preference of Geoffroy’s cat for
dense ground cover may serve to separate the two species
ecologically (Fig. 9).

Protected
species

area
may

where
occur

Figure 9. Distributionof Geoff roy’s cat (0. geoffroyr).
1. Specimen
taken near Tiraque (Cabrera1957); 2. Rio Boppi
VI (Bolivia); 3. Defensoresdel Chaco II; 4. Teniente Encisco II(Paraguay);
5. Quebradade 10s Cuervos
reserve (proposed);
6. Laguna de Castillos IV complex (Uruguay);
7. Rio Pilcomayo
II; 8. Fuerte EsperanzaI; 9. lber IV complex;
10. Campos delTuyu I complex;
11. El Payen I; 12. Sierra de la Ventana II;
13. Specimen
taken just east of Bariloche,
possible occurrencein Nahuel Huapi II complex (Melquist
1985, O.N. Herrera in litt.
1992); 14. Meseta de Somuncura
VIII; 15. Peninsula
de Valdes
VIII; 16. Perito Moreno
IV (Argentina);
17. Torres del Paine II*complex (Chile).

Population Status
Global: Category 4. Regional: Category 3. IUCN: notlisted. Throughout its range, Geoffroy’s cat has been
described as the most common of the small cats (Ximenez

130Part I: Species

Accounts.

Chapter

5. The Americas,

Puma

tor, and its adaptability probably helped it survive the late
Pleistocene extinctions of the other large North American
felids. Although it is a big cat, it is believed to be moreclosely related to the small cats, lacking the elastic hyoid
and enlarged vocal folds of the Pantherines (Hast 1989).
While it cannot roar, it is capable of a variety of vocalizations, and both sexes have a distinctive call, likened to a
woman’s scream, which is probably associated withcourtship (Young and Goldman 1946). Average weights
range from 53-72 kg for adult males and 34-48 kg for adultfemales (Anderson 1983, Pall et al. 1988), and males have
exceptionally weighed up to 120 kg (Banfield 1974).
Pumas tend to be larger away from the equator toward thepoles (McNab 197 1, Kurten 1973, Iriarte et al. 1990).
They have large feet and proportionally the longest hindlegs of the cat family (Gonyea 1976). The coat is plain(hence the Latin name covzcolor), which can vary in color
from silvery-grey
to tawny to reddish; as with thejaguarundi, coat color can be very different even between
siblings (P. Crawshaw, pers. comm.). Faint horizontal
stripes may occur on the upper forelegs (P. Jackson, pers.
comm.).
Melanism has been widely reported, andalbinism infrequently (Guggisberg 1975, Tinsley 1987,
Anon. 1989b). Young kittens are spotted, with blue eyes.
The known prey of pumas ranges from insects, birds,
and mice up to porcupine, capybara, pronghorn, wapiti,
bighorn sheep, and moose (reviewed by Guggisberg 1975,
Anderson 1983, Lindzey 1987, Iriarte et al. 1990, Hansen
1992). Large ungulates, particularly deer, are the puma’s
principal prey in North America. Studies conducted in
the region have found that ungulates make up an averageof 68% of the diet (Iriarte et al. 1990). Ackerman et al.
(1986) suggest that the energy requirements of femaleswith young are such that viable populations cannot exist inareas devoid of deer-size ungulates. For example, they
predicted that a resident female (based on studies in southem Utah) would kill a white-tailed deer every 16 days, andthat the interval would shrink to nine days when her kittens
were three months old, and to three days when the kittens
were nearly mature at 15 months.
However, in the southern parts of puma range, and particularly in the tropics, small to medium-sized prey appear
to be more important. Although the studies on which thisconclusion is based suffer from extremely small sample
sizes (Rabinowitz and Nottingham 1986, Emmons 1987,
Crawshaw and Quigley in prep.), Iriarte et al. (1990) suggest that the puma’s smaller body size in the tropics, andits low rate of predation on larger prey (such as tapirs),
are linked to interspecific competition with the jaguar (thelargest American felid). Pumas in the tropics are probably still capable of taking large prey, but Crawshaw andQuigley (in prep.), working on a ranch in the subtropicalBrazilian Pantanal, found that while pumas take calves ofdomestic cattle and sheep, adult cattle were taken only bybeech forest and matorral shrubland), and at 0.7 per
10 km2 over the entire shrub/grassland mosaic found in thepark (Johnson et al. in press).

fagus

Protection Status
Upgraded to CITES Appendix I in 1992. National legislation: fully protected across its range. Hunting prohibited:
Argentina, Bolivia, Brazil, Chile, Paraguay, Uruguay.
Domestic trade prohibited by all range states (Fuller et al.
1987, Govt. of Brazil).
Principal Threats
The species has been exploited commercially since theinternational cat skin trade boomed in the late 1960s withnearly 350,000 skins exported from Argentina alone
between 1976 and 1979 (Mares and Ojeda 1984). Tradevolumes remained high into the 1980s as trade in ocelot
pelts declined, averaging 55,000 per year between 19801984 (Broad 1987). Paraguay and Bolivia were the main
exporters (in contravention of national legislation) during
this time, although it is believed that the bulk of these skins
were smuggled in from Brazil and Argentina.
International trade has since declined-no significant trade has
been reported since 1988 (Govt. of Brazil 1992, WCMC
unpubl. data). Paraguay and Uruguay remain as important domestic markets for pelts (Brooks 1992, Ximenez in
Zitf. 1990). However, Walton (199 1) reports that mostpelts in trade today are derived from cats killed as pests
and livestock predators, and that commercial hunting as itexisted in the past has essentially ceased.
Geoffroy’s cats appear to tolerate moderate levels ofdeforestation (Koford 1973, Brooks 1992). So little is
known about the species’ ecological requirements that it
is at present impossible to judge the actual impact of hunting or habitat loss.
Action Planning
Project 99.

Puma, Puma concolor(Linnaeus, 1771)
Other Names
Cougar, mountain lion, catamount, panther (English);puma (French); Puma, Silberlowe (German); lean, leon
Colorado, lean de montafia (Spanish); onca vermelha, onca
parda su$uarana (Brazil); leon sabanero (Colombia); tigrouge (French Guiana); guasura, yagu-pyt (Guarani); cabcob (Mayan); leopard0 (Mexico); reditigri (Suriname).
Description
and Behavior (Plate 4)
The puma is an exceptionally successful generalist preda-

131Part I: Species

Accounts.

Chapter

5. The Americas,

Puma

jaguars. Still, even in Chile, where jaguars do not occur,
small prey is a key element of puma diet. Pumas in themountains of southern Chile eat 25 European hares forevery one guanaco (Johnson et al. in press). Iriarte et ~22.
(199 1) point out that the European hare was introduced toSouth America only 90 years ago, and its significance asa prey item for pumas demonstrates the flexibility of theirpredatory behavior. It is interesting that the puma occurs
in a variety of habitats and takes both large and small prey,similarly to the leopard in the Old World, while the jaguar,
like the tiger, is closely tied to well-watered forested environments and is capable of taking very large prey.
Radiotelemetry studies in North America and southern
Chile have found pumas to be primarily nocturnal and crepuscular, with activity peaks at dusk and dawn, and limited
diurnal activity (Van Dyke et al. 1986a, review by Hansen
1992, Johnson et al. in press). Males make scrapes inprominent locations, and especially along boundaries ofhome ranges. This behavior apparently advertises temporal presence (Hornocker 1969, Seidensticker et al. 1973).
Large kills are often covered with scraped-over vegetation and dirt, and pumas often remain in the vicinity,
returning frequently to feed. Seidensticker et al. (1973)

Age at independence: (W) 12- 18 months (Anderson 1983).
Age at sexual maturity: (W) both sexes 24 months, andfemales sometimes as early as 20 months (Lindzey 1987,

F. Lindzey unpubl. data), but time of first breeding probably depends on when a female is able to establish her ter-ritory (Homocker 1970, Seidensticker et al. 1973). Loganet al. (1986) found that females only entered the breeding
population at age 3-4 years in their hunted study populationin Wyoming. Females in stable populations rarely breedwith more than one male during es&us (Hemker 1982).
Recruitment rates: (W) 1.O- 1.3 kittens per breeding female(Alberta: Jalkotzy et al. 1992).
Interbirth interval: (W) can be one year (Robinette et al.
196 1), but more generally 18-24 months (Lindzey 1987).
Adult sex ratio: (W) most studies report 2: 1 female:maleratio of breeding adults, although other ratios, both higher
and lower, have been found (reviewed by Beier 1993).
Adult mortnlity rates: (W) Natural mortality appears to be
low, on the order of <5% (Homocker 1970, Currier et al.
1977, Ashman et al. 1983, Murphy 1983, Logan et al.

found that, in the winter, one puma fed from a carcass for1986). Mortality caused by sport hunting can be high, par-19 days. However,ticularly among adult and sub-adult males. Sport hunting

pumas rarely feed from carcasses ofthemselves have not killed (F.
Lindzey in litt. 1993).

in Alberta accounted for 63% of known mortality of radio-

animals which theycollared pumas (n=lO of 16) from 198 1- 1989, and 100%of sub-adult males (n=5) (Ross and Jalkotzy 1992).
Mortality caused by intraspecific conflict may be higher inboth populations which are hunted, where immigrantscompete to establish territories (Logan et al. 1986), and in
populations where food resources are relatively scarce,
such as semi-arid desert in New Mexico (Hornocker 1992,
Sweanor and Logan 1992).

Biology
Reproductive season: (W) Probably year-round (Sweanorand Logan 1992, Ross and Jalkotzy 1992), although mostbirths are reported to occur in the warmer months of AprilSeptember in the north of their range (Robinette et al.
1961, Eaton and Verlander 1977, Ashman et al. 1983,
Lindzey 1987). In the Torres de1 Paine National Park insouthern Chile, all known births (n=4) took place between
February-June (Johnson et al. in press).

Longevity: CW) probably 8- 10 (Hansen 1992), but up to
12- 13 years (Currier 1983), and a female puma on
Canada’s Vancouver island was killed by hunters when

she was at least 18 years old (M. Jalkotzy in litt. 1993); (C)Estrus: (C) 8 days.

up to 2 1 years (Hansen 1992).

Estrus cycle: (C) 23 days (Hansen 1992).
Gestation: (C & W) 91.9 * 4 days.

Habitat and Distribution
Pumas have a very broad latitudinal range encompassing

Litter size: (C & W) average 2.2-2.7 (Anderson 1983,
Currier 1983, Ross and Jalkotzy 1992); range l-6; possibly
only single kitten first litter (Lindzey 1987).

a diverse array of habitats (Fig. lo), from arid desert totropical rain forest to cold coniferous forest, from sea levelup to 5,800 m in the Andes (Redford and Eisenberg 1992).
While several studies have shown that habitat with dense
understory vegetation is preferred (Seidensticker et al.
1973, Logan and Irwin 1985, Laing 1988, Johnson et al. inpress), pumas can live in very open habitats with only a

Cub survival: (W) Hemker et al. (1986) estimated survival of cubs to dispersal at 67% in a non-hunted population in southern Utah, and suggested that cub survival
would be less in hunted populations, particularly if hunt-

ing seasons coincidedwith seasonal birth peaks.

minimum

However, Ross and Jalkotzy (1992) found 97% cub survival in a hunted population.

of vegetative cover (Lindzey 1987, Seiden-

sticker 1991 b). Pumas are occasionally reported fromareas of intensive agricultural cultivation, although such

132

Part I: Species

Unconfirmed

Accounts.

ChapterrecordFigure 10. Past and present
distributionof the puma (I? conco/o~$
1. Los Glaciares
II** (Argentina)+
Torres del Paine II* (Chile) complex; 2. Laguna San Rafael II* complex;
3. Vicente Perez Rosales II complex;
4. Lauca II* complex (Chile); 5. Los Alerces II, Lanin II + Nahuel Huapi II complex; 6. San Guillermo
IV* complex;
7. Los Andes I; 8. Peninsula
de Valdes VIII (Argentina);
9. lguazu II** (Argentina)+ Iguacu II** (Brazil) complex;
IO. Pacaas Novos II complex;
11. Amazonia
II complex;
12. Araguaia
II complex;
13. Lago Piratuba I (Brazil);
14. Defensoresdel Chaco II (Paraguay);
15. lsiboro Secure II; 16. Noel Kempff Mercado
II; 17. Manuripi Heath IVcomplex (Bolivia); 18. Manu II#; 19. Huascarn
II# (Peru); 20. Ecuador parks: Sangay II**, Cayambe-CocaI, andYasuni II complex; 21. ChiribiqueteII; 22. Sierra Nevada de Santa Marta II* complex (Colombia);
23. Serrania
dela Neblina II (Venezuela)+ Pica da Neblina II (Brazil) complex;
24. Canaima
II; 25. Aguaro-Guariquito
II (Venezuela); 26. Rio Platano II* (Honduras);
27. Calakmul
V* (Mexico)+ Maya IX* (Guatemala)complex;28. Big
II** complex;
Cypress
II; 29. Big Bend II*; 30. Grand Canyon
II ** complex; 31. Death Valley III; 32. Yosemite
33. Yellowstone
II# complex;
34. Olympic II# (U.S.); 35. Glacier II* (U.S.) + Water-ton Lakes II (Canada);
36. Jasper II complex; 37. Wells Gray II; 38. Tweedsmuir
II; 39. puma shot on Wrangell Island in 1989; 40. puma
sightings in the Kluane Lake region (Tischendorf
and Henderson
1993) (Canada);
41. Black Hills National
Forest/Custer
State Park complex;
42. Ozark/Ouachita/Mark
Twain National Forest complex; 43. Young malepuma captured
in 1991 in agricultural
region of southwest
Minnesota;
44. Puma shot near Lake Abitibi in 1992
(Tischendorf
and Henderson
1993); 45 tracks and scat found in east-central
New Brunswick(Cumberland
1993).
133

5. The Americas,

Puma

Part I: Species

Accounts.

Chapter

5. The Americas,

Puma(Tischendorf and Henderson 1993). The puma has faredmuch better in the less populated western U.S., and withchanges in management status from “varmint” (vermin)
to game animal, numbers there appear to be increasing(reviewed by Beier 1991, Seidensticker and Lumpkin1992). Population estimates by state wildlife authorities
indicate that pumas in the western U.S. probably number
over 10,000 (Tischendorf 199 1).
Anderson (1983) summarized information on densitiesof puma populations in North America. For those populations that were studied for at least 12 months or overtwo consecutive winters, densities ranged from 0.5 to 4.9
individuals/l00
km2 (studies carried out only in North
America). The lowest densities have been reported fromarid regions (Hemker et al. 1984, Sweanor 1990). A protected population in Patagonia (50” S) was found to havean average density of 7 animals/100 km2, among the highest densities so far documented, and presumably the resultof protection and an abundant prey base (Iriarte et al. 199 1,
Johnson et al. in press). Crawshaw and Quigley (unpubl.
data) estimated density at 4.4 individuals per 100 km2 ona cattle ranch in the Brazilian Pantanal. Based on a number of North American studies, Shaw (1989) concludes
that adult resident carrying capacity is of the order of 26-52
km2 per individual. Densities have not been calculated
for pumas inhabiting tropical forests.
Lindzey (1987) summarized the results of North
American studies of puma home ranges: average range
size varied from 32- 1,03 1 km? Male home range size
tended to be at least several hundred km2, while mostfemale ranges were less than 100 km2. Male home ranges
typically encompass those of several slightly overlapping
resident females; resident male ranges only occasionallyoverlap. However, one study in central California’s
Diablo Mountains found that male ranges overlappedwhile those of females did not (Hopkins 1989). The
largest home ranges have been found in arid environments(McBride 1976), while the smallest documented homeranges appear to be in areas where the major prey is nonmigratory (Sitton 1977). Pumas living in mountains thatreceive heavy snowfall tend to shift their ranges downward
in elevation, following seasonal movements of ungulates(Seidensticker et al. 1973, Ashman et al. 1983, Hemker etal. 1984).

animals are likely to be transient (TischendorfandHenderson 1993).
The puma’s historical distribution included every major
habitat type in the Americas up to the boreal forests of the
far north, but pumas have been essentially eliminated fromeastern North America. Severe reduction of native ungulate populations through hunting and forest clearance during the nineteenth century, coupled with direct persecutionof the puma, are the probable causes (Wright 1959). Deer
have since multiplied and spread, and the puma is now
found in areas colonized by deer which were outside its
historical range, such as the Great Basin Desert in thewestern U.S. (Berger and Wehausen 1991).
Population Status
Global: Category 5a(A). Regional: Category 4(A). IUCN:
eastern cougar (F.c. cougar) and Florida panther (F.c.
coryi; see Box 2) Endangered in North America.
In
Central and South America, the puma still occurs throughout much of its historical range. However, focused studies
have only been carried out in North America.
In Canada, pumas have been extirpated from most of
their former range, and the main population is now foundin southwestern British Columbia, where they are estimated to number 3,500~5,000 (Hummel 1990). In adjacent habitat in Alberta, Jalkotzy et al. (1992) estimated apopulation of 685 pumas, with 93% living outside nationalparks. There are occasional reports of pumas in the far
south of the Northwest Territories (K. Poole in Zitt. 1993),
and in Saskatchewan, Manitoba, Ontario, Quebec, New
Brunswick, and Nova Scotia (Anon. 1989b). Presence in aremote forested area of east-central New Brunswick wasrecently confirmed by the finding of a set of tracks and scat
(Cumberland 1993). In Manitoba, on the other hand, while
puma reports are made to the government at a rate of lo-50
per year (Johnson 1990), the only field survey produced noevidence of puma presence (M. Jalkotzy in Zitt. 1993). An
Eastern Panther Recovery Team has been formed by the
Canadian Wildlife Service (Tischendorf 1992a).
As in Canada, the puma was essentially eliminatedfrom most of the eastern U.S. within 200 years following
European colonization (Wright 1959). The only easternstate where the puma is unequivocally known to persist is
Florida (see Box 2). In the northeastern U.S., hundreds ofsighting reports have been investigated and compiled(Wright 1972, Anon. 1989b, Tischendorf 1992b), but theexistence of actual remnant populations has not been verified. Several networks have been established to further
investigate the species status in the region, including theEastern Puma Research Network and the Friends of theEastern Panther (Tischendorf 1992a). Pumas are rare in
the central plains region, with few resident populations

Protection
StatusCITES Appendix II; eastern and Central American subspecies (F. c.coryi, costaricensis, and cougar) Appendix
I. National legislation: protected over much of its range.
Hunting prohibited: Argentina, Brazil, Bolivia, Chile,
Colombia, Costa Rica, French Guiana, Guatemala,Honduras, Nicaragua, Panama, Paraguay, Suriname,134Part I: Species

Venezuela, Uruguay. Hunting regulated: Canada, Mexico, Peru, United States. No legal protection: Ecuador, El
Salvador, Guyana (Fuller et al. 1987, R. Hoogesteijn inlitt. 1993).
There is good information available about the protection status of the puma on a regional level in North
America. The easterncougar is protectedin the Canadianprovinces of Nova Scotia, New Brunswick, and Ontario(Macey 1979). Elsewherein Canada,pumasareprotectedin Manitoba, Saskatchewan,and Yukon, but not at all in
the Northwest Territories. Hunting is regulatedin Alberta
and British Columbia. In the U.S., the easterncougar isfully protected under the EndangeredSpeciesAct. In thewest, pumasare fully protected only in South Dakota andCalifornia. Hunting is regulated in Arizona, Colorado,
Idaho, Montana, Nevada, New Mexico, Oklahoma, Oregon, Utah, Washington, and Wyoming. Pumas are notlegally protected in Texas (Tischendorf 1991).
In California, which holds one of North America’s
largestpumapopulations,voters in 1990narrowly approved an initiative which directed the state to prohibit sport
hunting of pumas (formerly permitted), and to allocate
U.S. $30 million a year for the next 30 years toward provision of habitat for pumasand other threatened species(West 1991).

Accounts.

Chapter

5. The Americas,

Puma

Box 2
The Florida Panther,
Puma concolor coryi (Bangs, 1899)
A Worst-Case Scenario
The problems involved when wild cats are reduced toa minimal level and they live in a human-dominatedarea, are well illustrated by the case of the puma subspecies known as the Florida panther. The threatsfacing the panther are numerous and complex, andare indicative of the likely fate of big cats around settled areas if preventive conservation measures are
not taken. A remarkable,
broad-based
effort has
been mounted to save the Florida panther, and thecomplexity, difficulties, and costs involved demonstrate the importanceof taking action to conservecat populations before they become seriously threatened.
The Florida panther was formerly found throughout
the southeastern United States, but had disappeared
from most of its range by the late 1920s (Lowery
1936, Young and Goldman 1946, Tinsley 1970).
Florida was one of the first states to offer any legal
protection to the panther (in the 195Os), and it is now
home to the only known puma population in eastern
North America, consisting of just 30-W adult animals(Jordan 1994) confined to fragmented
patches ofhabitat (Fig. I I).

Principal Threats
Across the Americas, ranchers are likely to continue toview pumasasa threat to their livestock and to attempt toelimatethem. Pumasarevulnerablebecausethey return totheir kills, which can be poisoned,andbecausethey take totreeswhen huntedby dogs. Shaw(1977) found that calvesof lessthan one year, weighing generally under 90 kg,
were most frequently taken in Arizona. Crawshaw andQuigley (in prep.) found similar results on a Brazilian
ranch, and Yafiez et ~2.(1986) observedthat pumaswere
significant predators of sheepon ranchesin Chile. See
Part II, Chapter2 for further discussionof livestock depredation by pumasand ways to control it.
With legal protection, pumasnow occur very close tosettled areas throughout western North America, andattackson people,while infrequent comparedto other hazards from animals or nature (such as lightning strikes),
have increased(Beier 1991). However, there seemsto begenuinepublic supportfor the puma’spresencein mostof
North America despitethe dangers,a major changein public perceptionover the pastfew decades(Seidenstickerand
Lumpkin 1992,Jalkotzy et al. 1992).

One sub-population(Everglades
National Park)
became extinct in 1991, when the last two females
known to live in the area died (Hansen 1992)-theEverglades are only occasionally frequented now by
one male (D. Jordan, pers. comm. 1994). Everglades’ panthers had been previously analyzed
genetically and were found not to be pure coryi: they
carried genes from pumas of partial South American
descent released in the Everglades in the late 1950s
to early 1960s (O’Brien ef al. 1990). In comparison
with the main Florida panther population(Big
Cypress swamp ecosystem), this introgressionof
new genetic material could be advantageous, as the
Big Cypress animals have a number of physiologicalimpairments
which can be ascribed to inbreeding
depression,caused by population
isolation anddecline. These include a high (95%) abnormal sperm
count among males, cryptorchidism(one or two
undescended
testicles), heart murmurs, and vaginal
fibropapillomas (possibly impeding reproductive success) .
Other health problems
affecting the population
include exposure to domestic animal-borne diseases,
such as feline panleukopenia,
rabies,felineHIV and parvovirus(genetic homogeneity
may

Action Planning
Project 6 and 100-103.

Continued on next page

135

Part I: Species

Accounts,

Chapter

5. The Americas,

GEORGIA

c“\,

Puma

less productive Jand that has been given over to the
public trust. Studies have shown that panthers thatprimarily inhabit private lands are in better physicalcondition and have a higher rate of reproductive success than individuals on public lands, possibly due
to greater prey availability and less disturbance fromhunters (Maehr 1990).

v

A population
viability analysis conductedby theIUCNISSC’s Captive Breeding Specialist Group concluded that the Florida panther could become extinct
within 25-40 years under prevailing demographic andgenetic conditions (Seal et al. 1989).
There are a number of lessons to be drawn from thecase of the Florida panther.
The response of theAmerican conservation authorities has been impressive and has involved creative management
techniques.
Followinga private initiative in 1976, a
Florida Panther Recovery Team was establishedwith representativesof various Federal and Stateagencies and other experts and a full-time Florida
Panther Coordinator.
The first task was to find out if
panthers survived and their location. A radio telemetry program was initiated amid public controversy
which intensified when a panther died after being
darted with a tranquilizer.
However, the program
continued with official approval. In 1986, a Florida
Panther Interagency Committee was established andwent on to work with private landowners to develop
a program to preserve vital corridors and strategic
parcels of habitat, with priority initially allocated to
3,752 km2 of privately-owned
land (Maehr 1990,
Logan et al. 1993).

.. . . . . . . . . . . . .
..
\ ..
Exrstrng
Conservation
Lands

Figure 11. Distributionof the Florida
panther(I? concolor
cov$
Range based on radio-instrumented
segment of the population(19 panthers/May
1994).
Source: Dennis B. Jordan, Florida Panther Coordinator,
U.S. Fish and Wildlife Service.

impede immune response), and events which may
be due to poor nutrition, such as anemia and parasitic infestation (Roelke et al. 1993). Between 19790
1991, road kills accounted
for half of all known
panther deaths (11 out of 22: Maehr ef a/. 1991 a).
Very high levels of mercury were found in two of thedead Everglades
panthers, possibly from eatingracoons which had eaten contaminated fish (Jordan
1990, Hansen 1992).

However, government response has still been hampered by bureaucracy.
A former member of thePanther Advisory Council has documented the difficulties which arose in inter-agency cooperation anddeclared that problems with the Florida panther program “focus attention on a poorly understood impediment of the recovery of endangered
species in the
United States: a land management bureaucracy thatwill not acknowledge
the novel demands of small
population biology-therequisite discipline” (Alvarez1993).

These problems have taken on enormous dimensions because little habitat remains to support larger
numbers of panthers. What is left of the Florida panther’s habitat is fragmented by agriculture and settlements and crisscrossedby roads. Some of themost strategically located forested areas are privately
owned, and some owners are hostile to the pantherrecovery efforts for fear of losing land-use optionssuch as citrus growing. Of approximately 12,555 km2
of occupied panther range in south Florida, only 47%
is in state and federal ownership (Logan Hal. 1993).
None of the radio-collared
panthers has restricted its
movements solely to public lands. Moreover, as is
the case across much of the world, it is marginal or

The impact of a major highway which cuts throughthe heart of panther country is being lessened by theconstruction of fences and 36 underpasses,
undertaken at a cost of $30 million (Harpster 1990); panthers have been using the underpasses(Humphrey
ef a/. 1991). Emphasis is being given to ensuring
adequate densities of white-tailed deer and wild hogs
by control of sport hunting and other recreational
activities on public lands. Wild panthers have been
vaccinated against several diseases. Seven kittens
have been removed from the wild and placed in aContinued

136

on next pagePart I: Species

Accounts.

Chapter

5. The Americas,

OcelotOcelot, Leopardus paralis(Linnaeus, 1758)

captive-breeding
program to build up a reserve population to supplement panthers in the wild (Jordan
1991). The panther has been designated the official
Florida state animal.
Public support has beenremarkably strong, and interest has been generated
by the Florida Power and Light Co.‘s publication anddistribution of a popularized but thorough discussionof the Florida Panther Recovery Plan (USFWS 1987,
Van Meter 1988).

Other Names
Ocelot (French); Ozelot (German); tigrillo, ocelote, gato
onza (Spanish); tirica (Argentina); gato onza (Argentina,
Bolivia, Peru); chivi-guazu (Argentina, Paraguay);
cunaguaro(Argentina, Venezuela); tigrezillo, gato bueno(Bolivia); maracaju-acu, gato mourisco, jaguatirica(Brazil); gato maracaja (Brazil, Paraguay); maracaya,
maracaja(Colombia); manigordo(CostaRica, Nicaragua,
Panama,Venezuela); chat tig (French Guiana); yagua-tirica (Guarani); zac-xicin (Mayan); gato tigre, tigre Chico(Panama);pumillo, tigrillo (Peru); hetigrikati (Suriname).

The government
has set the long-termgoal ofachieving three viable, self-sustaining
populations
within the panther’s historic range (Jordan 1993).
Given the unlikelihood of the current population thriving and expanding,reintroductionis the government’s preferred management
strategy, and it iscurrently carrying out its second reintroduction
feasibility study. In the first attempt, seven wild-caught
Texas panthers
were releasedin the Osceola
National Forest, in northern Florida on the boundarywith Georgia. While there was evidence of successful land tenure, all were recaptured
earlier thanplanned due to conflicts with humans-one,forexample, climbed up a tree in a Jacksonville backyard (Bolgiano 1991). The latest attempt involves 10
Texas panthers-three
captive-bred and seven wildcaught adults-whichwere radio-collared
andreleased in the same area in February 1993 (Belden
and McCown 1993: Fig. 11). It is interesting that,after four months of monitoring, the captive-raised
panthers (which had been given pre-release training
in catching live prey) appeared to have settled down
more quickly than the wild-caught cats, using smallerareas and more frequently capturing prey (Belden
and McCown 1993). However, the same problems
which occurred in the last study are happening
again. Three cats (two wild-caught,
one captiveraised) have been recaptured,and others (wildcaught) relocated after people complained that thecats threatened life or property (Belden and McCown
1993, Jordan 1994). The recapture of the captiveraised male occurred after he was seen killing ahouse cat (D. Jordan, pers. comm. 1994). This provides a good example of the difficulties of reintroducing and conserving big cats near people.

Description and Behavior (Plate 5)
The ocelot is the bestknown smallcat of the Americas, as
far as the public is concerned, largely because of thebeauty of its coat. Its pelageis short and close(lessplush
than the similarly patterned margay), and marked withboth rosettesand spotswhich tend to run in parallel chains
along the sides of the body. It is also the best-studied(Navarro 1985,Tewes 1986, Emmons1987, Ludlow andSunquist 1987, Emmons 1988, Crawshaw and Quigley
1989,Emmonset al. 1989,Konecny 1989,Sunquistet al.
1989; P. Crawshaw, M. Tewes, pers. comm.). Average
weight of adult malesis 10 (n=8: Mondolfi 1986) to 11.5
kg (n=8: Enders 1935, Emmons 1988, Sunquist et al.
1989), and females 8.8 (n=5: Mondolfi 1986) to 9.4 kg(n=l 1: Husson 1978, Emmons 1988, Crawshaw andQuigley 1989,Konecny 1989, Sunquistet al. 1989).
Thesestudiesfound terrestrial andnocturnal rodentsto
be the mainstayof the ocelot’sdiet. Most frequently taken
prey specieswere thoseof relatively high abundance,and
included canemice and marsh,spiny, and rice rats (Bisbal
1986,Ludlow and Sunquist 1987,Emmons 1988), opossumsand armadillos (Konecny 1989). Ocelots will also
take larger prey, including lesser anteaters (Mondolfi
1986,Konecny 1989),red bracket deer (Konecny 1989),
squirrel monkeys (Emmons1988), and land tortoises(thelegs of a very young tortoise were found in an ocelot’s
stomach: Mondolfi 1986). However, most prey taken
weighslessthan l-3% of anocelot’s body weight (Ludlow
andSunquist 1987,Emmons1987);for larger prey species(such as paca and agouti), juveniles are typically taken(Emmons 1987). Ocelots also vary their hunting behavior to take advantage of seasonalchangesin prey abundance, such as spawning fish (Emmons 1988) and landcrabs (Ludlow and Sunquist 1987) in the wet season.
Basedon scatanalysis,Sunquistet al. (1989) found ocelot
diet to consistof 65% smallrodents, 18%reptiles (mostly
iguanas), 7% crustaceans and fish, 6% medium-sized

The Florida Panther Interagency
Committee
isattempting to deal with the genetic problems of thepanther by giving conceptual approval to restoringhistoric gene flow (USFWS 1993), either througheventually facilitating connectivity between the south
Florida panthers and reintroduced
Texas panthers,or through interactive outbreeding with captive animals. These plans are complicated by the fact that
the U.S. Endangered
Species Act may not extend
protection to progeny resulting from intercross breeding. A policy determination on this matter is presently
under development.

137

Par? I: Species

Accounts.

Chapter

5. The Americas,

Ocelotdense vegetative cover is the common factor linking thevarious habitat types where ocelots are found (Navarro
1985, Tewes 1986, Ludlow and Sunquist 1987). Emmons(1988) suggests that ocelot microdistribution
is morepatchy than would
uld be expected by its wide geographicalrange (Fig. 12),1;; depending upon an abundant rodent preybase and good ground cover. Ocelots generally venture
into open areas only on moonless nights or cloudy days
(Ludlow and Sunquist 1987, Emmons et al. 1989).
The ocelot’s known range was extended 350 km southwards when its presence was confirmed in Uruguay(Ximenez 1988). At the northern end of its range, only
two significant ocelot populations are believed to persist in
the southeastern corner of Texas (Tewes and Everett1986). Ocelots have been extirpated from Arkansas,
Louisiana, eastern Texas, and Arizona, although individuals may occasionally cross into Arizona from Mexico.

mammals, and 4% birds. Similarly, Emmons (1988)
found ocelot diet to consist of 66% small mammals, 5%large rodents, 5% bats and arboreal mammals, 11% birds,
12% reptiles and 2% fish.
Ocelots are strongly nocturnal, resting in trees or densebush in the daytime; some resting spots are used repeatedly(Emmons 1988), including by different ocelots of the samesex (Ludlow and Sunquist 1987). Ocelots are generally
active for more than half of each 24-hour period. Mean
daily travel distances range from 1.8-7.6 km, with males
travelling up to twice as far as females (Ludlow andSunquist 1987, Emmons 1988, Konecny 1989). Emmons(1988) estimated ocelot hunting success at 0.9 prey captured per km walked, or one prey captured for every 3.1
hours of travel. Her observations of ocelot hunting behavior suggest that they can follow prey odor trails.
Biology
Reproductive season: (W) Probably year-round, with
autumn breeding peaks noted in Texas (Tewes 1986) andMexico (Leopold 1959); October-January
peaks also
reported from Paraguay (Rengger 1830) and northeastern
Argentina (Crespo 1982).

Population Status
Global: Category 5a. Regional: Category 4. IUCN: notlisted. The ocelot was the spotted cat most heavily
exploited by the fur trade from the early 1960s to the mid1970s: Gieteling (1972) estimated that as many as 200,000
animals were taken every year. From 1976 to 1983, net
international trade in skins fell to an average of 24,600
pelts annually, and effectively ceased in the late 1980s(Broad 1987, WCMC unpubl. data) (see Part II Chapter 4).
In the early 1980s Mondolfi (1986) reported that commercial hunting had depleted formerly abundant ocelot
populations
in Venezuela, but more recently R.
Hoogesteijn (in Zitt. 199 1) noted that hunting pressure is
now greatly reduced, and there are signs of recolonization
and recovery. L. Emmons (in Zitt. 1993) notes that, even at
the lowest density estimates (one animal per 5 km2), there
would be approximately 800,000 ocelots in forested SouthAmerica alone, and suggests that true numbers are probably 1.5-3 million.
The ocelot is one of the few small small cats for which
spacing and abundance have been studied in several different habitat types.

Estrus: (C) 4.63 k 0.63 days (n=6).
Estrus cycle:(C) 25.11 t 4.33 days (n=9: Mellen 1989).
Gestation: (C) 79-85 days (Mondolfi1986).

Litter size: (C) 1.64 t 0.21 (n=28: Mellen 1989); range l3, mode 1 (Cisin 1967).
Age at independence: (W) not clear, approximately one
year, after which sub-adults appear to be tolerated within
adult ranges for up to a year (Ludlow and Sunquist 1987,
Emmons 1988, Crawshaw and Quigley 1989).
Age atfirst reproduction: (C) females 18-22 months, onsetof spermatogenesis in males at about 2.5 years (Mondolfi1986); (W) probably related to territory acquisition (L.
Emmons in Zitt. 1993).
Interbirth1988).

interval,(W) possibly

two years (Emmons

Lowland rain .forest

Longevity: (W) 7- 10 years; (C) up to 20 years (Sunquist
1992). ”

1. Manu National Park (Peru): Home ranges of two adultfemales were 1.6 and 2.5 km2; home ranges of two
adult males were 5.9 and 8.1 km? Resident animals,
particularly males, often patrolled the perimeter of theirterritories, travelling quickly (8.8-10.4 km/hr) and seldom pausing. Density was high, estimated at four resident ocelots per 5 km’ (Emmons 1988).

Habitat and Distribution
The ocelot is found in every country south of the UnitedStates, except Chile, and occupies a wide spectrum of
habitats, including mangrove forests and coastal marshes,
Savannah grasslands and pastures, thorn scrub, and tropicalforest of all types (primary, secondary; evergreen, seasonal, and montane-although
it typically occurs at elevations below 1,200 m [Mondolfi 1986, Bisbal 1989,
Eisenberg 19901). Availability of sufficient amounts of

2. Iguacu National Park (Brazil): From a radiotelemetry
study in progress, P. Crawshaw (in Zitt. 1993) reports
average home range size for six adult ocelots (2 M, 4 F)
as 11.3 km”.

138Part I: Species

Accounts.

,::.

:

Species

:

range

l
.
.
m
.
.

q
q
q
.

Protected
area where
species occurs

..
.9
i

l.
.

3(c

Protected
area where
species may occur.

Protected
conserve\

area over 3,000 Km* in size, possibly sufficienttoa minimum viable populationof over 1,000 breeding

.

..
l.
.1.

i

adultsFigure 12. Distributionof the ocelot (L. par&/k).
1. Laguna Atascosa
IV (Texas, U.S.); 2. Montes Azules II* (Mexico);
3. Calakmul
V* (Mexico)+ MayaIX* (Guatemala)complex; 4. Rio Platano II* (Honduras);
5. La Amistad II* (Costa Rica and Panama);
6. Darien II# (Panama);
7. Sierra Nevada de Santa Marta II*; 8. Sierra de la Macarena
II complex;
9. Chiribiquete
II (Colombia);
10. Perij II; 11. Serrania
de la Neblina II (Venezuela)
and Pica da Neblina
II (Brazil) complex;
12. Canaima
II; 13. Aguaro-Guariquito
II (Venezuela);
14. Trinity Hill IV (Trinidad
and Tobago); 15. Yasuni II* (Ecuador);
16. Cerros de Amotape
II complex;
17. Pacaya-Samiria
VIII;
18. Yanachaga
Chemillen
II; 19. Manu II# (Peru); 20. Cabo Orange II complex; 21. Jau II complex;
22. Amazonia(Tapajos)
II complex;
23. Araguaia
II complex;
24. Serra da Capivara
II; 25. Pantanal
Matogrossense
II; 26. Emas II; 27. Serra dos Orgaos II (Brazil); 28. Iguacu II** (Brazil) + Iguazu II**
(Argentina)
complex;
29. Sao Joaqim II (Brazil); 30. Bajaga da Pema and Meseta de Artigas reserves(proposed:
Uruguay);
31. Baritu II; 32. Fuerte Esperanza
I; 33. Agua Dulce Provincial
Reserve;
34. lber IV (Argentina);
35. TinfunqueII; 36. Defensoresdel Chaco II (Paraguay);
37. Beni I;
38. Manuripi Heath IV complex (Bolivia).

139

Chapter

5. The Americas,

OcelotPar? I: Species

Accounts.

Chapter

5. The Americas,

Bobcatpatches near human settlements (Koford 1976, R.
Hoogesteijn in litt. 199 1). Recent studies, however, depict
a more specialized animal operating under rather harshenvironmental constraints. Ocelots have a small average
litter size, and one of the longest gestations and slowest
growth rates among small felids (Fagen and Wiley 1978).
Emmons (1988) points out that these reproductive parameters suggest adaptation to low expected rates of energy
acquisition, a conclusion supported by findings that
ocelots spend long portions of each day hunting. One lactating female increased her daily activity by a maximum of
133% after birth of her litter, but still lost her young to starvation after six weeks-despitehigh diversity and abundance of small prey in the study area (Emmons 1988).
Emmons (1988) and Sunquist (1992) compare ocelots
to bobcats, which have a higher reproductive capacity,although both species are of equivalent weight. The maindifference is in their staple prey: for bobcats it is lagomorphs (about 10% of bobcat body weight), while for
ocelots it is a variety of small rodents. Emmons (1988)
suggests that ocelots may not be able to reproduce whereprey density is reduced. The two authors differ, however,on the potential impact of direct human hunting pressure.
Sunquist (1992) suggests that, given the ocelot’s low
recruitment rate, offtake of more than 2-3% of a population per year is likely to be unsustainable and result in
declines. Emmons (in litt. 1993), on the other hand,
believes that the ocelot populations have proved resilient
to harvest because of their social organization. A significant proportion of a population consists of non-breedingtransients (Emmons 1988), and thus loss of 2-3% of the
population would be unlikely to reduce breeding activity
substantially.

3. Cockscomb Basin Wildlife Sanctuary (Belize): Home
range of one adult female was quite large, 14.3 km2.
A sub-adult male had a home range of 3 1.2 km2. Mostof their territories consisted of secondary forest(Konecny 1989).
Seasonally flooded Savannah woodland
Venezuelan llanos: Home ranges of two adult males
were 9.3 and 11.1 km2. Mean home range for six adultfemales was 3.4 km2 (range 1.7-6.8). Density was estimated at two resident ocelots per 5 km2 (Ludlow andSunquist 1987).
Brazilian Pantanal: Home ranges (six months only) of
two adult females were 0.8 and 1.5 km2 (Crawshaw andQuigley 1989).
semi-arid woodland and scrub
6. Southern Texas (U.S.): In dense brush and oak forest
mosaic, an adult male maintained a home range of 3.5
km2 7and a female of 2.1 km2 (Navarro 1985). Working
in chaparral habitat, Tewes (1986) found a mean homerange for five males of 12.3 km2 (range 4.6-17.7) andfor three females of 7.0 km2 (range 4.9-9.9).
Protection Status
Upgraded to CITES Appendix I in 1989. National legislation: protected over most of its range. Hunting prohibited: Argentina, Brazil, Bolivia, Colombia, Costa Rica,
FrenchGuiana, Guatemala,Honduras,
Mexico,Nicaragua, Panama, Paraguay, Suriname, Trinidad,
United States, Uruguay, Venezuela. Hunting regulated:
Peru. No legal protection: Ecuador, El Salvador, Guyana(Fuller et al. 1987).

Action Planning
Projects 96 and 97.

Occurrencein Protected Areas
Adjusting for overlapping generations (females) and thefact that males typically mate with more than one female,
Ludlow and Sunquist (1987) calculated that a populationof 1,334 adult ocelots is required to realize an effective
population of 500 ocelots. They used a mean density of
0.38 adult ocelots per km2, as found for the Venezuelan
llanos, to derive a figure of 3,5 10 km2 required to support
a population of a size sufficient to minimize loss of genetic
diversity. Protected areas over 3,000 km2 are marked inFig. 12 with a square; other important protected areas arealso noted.

Bobcat, Lynx rufus(Schreber, 1776)
Other Names
Lynx roux, chat sauvage (French); Rotluchs, Luchskatze(German); lince, lince rojo, gato mantes (Spanish).
Description
and Behavior (Plate 6)
The bobcat, named for its short tail (TL=13-20 cm; 1316% of head-body length: Hall 198 l), is a medium-sized
cat with a ruff of fur edging the sides of the face. The average weight of adult males ranges from 8.9- 13.3 kg, andfemales from 5.8-9.2 kg (Anderson 1987). The largest
verifiable weight recorded is 17.6 kg for an adult male

Principal Threats
The ocelot has been labelled “one of the most successful
forms of mammalian life in the Amazon region”(Grimwood1969). It is tolerant of disturbed habitat(Vaughan 1983, Tello 1986b), and persists in wooded

140

Part I: Species

Accounts.

Chapter

5. The Americas,

Bobcatnight, but studies have consistently found crepuscular(dawn and dusk) activity peaks, a pattern based on theactivity patterns of major lagomorph and rodent preyspecies (reviewed by Anderson 1987, Rolley 1987).

from Minnesota (Berg 1979). Average adult weights andcranial measurements (Samson 1979) indicate that bobcats
are larger in the north of their range (Anderson 1987);
larger bobcats also tend to occur in more open habitats,with smaller bobcats in forested habitat (Read 1981). Pelt
coloration has been variously described as light grey, yellowish brown, buff, brown and reddish brown. Bobcats
are always spotted to some extent, with some individuals
patterned only on the undersides, others with spots extending up the sides onto the chest and back. Both melanistic
and albinistic specimens have been reported (Schantz
1939, Ulmer 1941, Young 1958)-melanism
has beenfound only in Florida (Regan and Maehr 1990). The bobcat may be distinguished from the similar-looking Canada
lynx by its shorter hind legs, smaller feet, and shorter ear
tufts. The bobcat’s tail is black only on the top, whereas
the lynx has black all around the tip.
However, as with the lynx, lagomorphs are the bobcat’s
year-round dietary staple-cottontail
rabbits in the southof their range, and snowshoe hares in the north (Maehr andBrady 1986, Anderson 1987, Rolley 1987). Unlike thespecialist lynx, however, the bobcat is a generalist and,depending on the locality, rodents also make an important
contribution to its diet (Young 1958). In the southerncentral Plains and the southern United States, cotton rats
are the primary food item (Kight 1962, Beasom and Moore
1977, Miller and Speake 1978, McCord and Cardoza
1982, Leopold and Krausman 1986, Maehr and Brady1986). Wood rats and kangaroo rats are important in thesouthwestern U.S. (Leach and Frazier 1953, Gashwiler etal. 1960, Small 1971, Jones and Smith 1979). In western
Washington state, Knick et al. (1984) reported a 42%
occurrence of mountain beaver in the diet. In Texas, peccaries are also taken (Leopold and Krausman 1986).
Bobcats also take birds (Leach and Frazier 1953, Beasomand Moore 1977, Miller and Speake 1978, Bailey 1979)and bats (Wroe and Wroe 1982).
Despite their small size, bobcats can be effective predators of large adult ungulates, especially deer, generally
killed when resting (Matson 1948, Young 3958). Several
studies have found that males tend to consume larger preythan females (Fritts and Sealander 1978a, Sweeney 1978,
Litvaitis et al. 1984), and that juveniles consume proportionally more rodents than adults (Whittle 1979, Toweill1982). Ungulates can be an important winter food sourcefor northern bobcats, when snow depth increases their vulnerability to predation (Erickson
1955, Fritts andSealander 1978a, Miller and Speake 1978, May 198 1,
Litvaitis et al. 1986, Koehler and Hornocker1989).
Young fawns are also particularly vulnerable (Beale andSmith 1973, Trainer 1975). Bobcats will also scavenge
ungulate carcasses killed by other predators (Koehler andHornocker 1991).
Bobcats may be active during all hours of the day andBiology
Reproductive season: (W) Breeding peak late winter(February-March),birth peak early spring (April-May),but in the south of their range litters have been recorded
from every month of the year (Anderson 1987, Rolley1987).
Estrus: (C & W) 5-10 days.
Estrus cycle: (C & W) approx. 44 days (Crowe 1975a,
Mehrer 1975). Bobcats may ovulate spontaneously rather
than by induction (Duke 1949, Crowe 1975a, Fritts andSealander 1978b).
Gestation: (C& W) mode 62, range 50-70 days (McCord
and Cardoza 1982).
Litter size: (C& W) average 2.6-2.8, range l-8. Younger
females produce consistently smaller litter sizes than older
adults (review by Anderson 1987).
Pregnancy rates: (W) as with litter size, probably density-dependent, and generally decline at times of low preydensity (Rolley 1983) or high bobcat population density(Lembeck and Gould 1979). Yearling pregnancy rates
have ranged from 26-46% (Bailey 1979, Parker and Smith
1983, Rolley 1985), while adult (>3 years) pregnancy rates
are much higher at 73-100% (Bailey 1979, Lembeck and
Gould 1979, Miller 1980, Parker and Smith 1983).
Interbirth interval: (W) generally one year (McCord andCardoza 1982, Anderson 1987, Rolley 1987).
Age at sexual maturity: (W) Females are physiologicallymature at 9-12 months (Crowe 1975a, Fritts and Sealander
1978b), but probably do not breed until their second year
(Pollack 1950, Fritts 1973, Crowe 1975a, Sweeney 1978).
The onset of breeding activity may be delayed during preyshortages (Rolley 1985) or in high density bobcat populations (Zezulak and Schwab 1979, Miller 1980). Onset ofspermatogenesis in males at 1.5 years (Fritts and Sealander
1978a).
Age at independence: (W) about one year, dispersal occurs
during peak of adult breeding activity (Crowe 1975b,
Bailey 198 1, Griffith and Fendley 1986, Kitchings andStory 1984).
Mortality: (W) From life history tables, Crowe (1975b)
estimated annual average juvenile mortality at 26% inWyoming from 1948-1973 (range 18-7 1%). Mortality isstrongly correlated with prey availability (Bailey 1974,
Knick 1990). In protected populations with an abundant141

Part I: Species

Accounts.

Chapter

5. The Americas,

BobcatUnited States, where they range through a wide variety ofhabitats, including boreal coniferous and mixed forests inthe north, bottomland hardwood forest and coastal swamp
in the southeast, and desert and scrubland in the southwest (Fig. 13). In the west, they have been trapped at elevations up to 2,575 m (Govt. of U.S. 1983a). Only large,
intensively cultivated areas appear to be unsuitable habitat.
Areas with dense understory vegetation and high prey density are most intensively selected by bobcats (Rolley1987). In Mexico, bobcats are found in dry scrubland andforests of pine and oak, principally in the mountainous
northern and central parts of the country, and not in thetropical south (Woloszyn and Woloszyn 1982, Gonzlez
and Lea1 1984).
Southern Canada represents the northern limit of bobcat
range. Bobcat feet are smaller than those of the lynx andlack the large furry pads (Banfield 1974). Parker et al.
(1983) found that the larger surface area of lynx paws supports twice the weight that of the bobcat can in the snow:
this, combined with the bobcat’s shorter legs, makes travel
in deep snow difficult, and is thus a significant limiting fac-prey base, natural adult mortality is low, estimated at 3%in Idaho (Crowe 1975b based on Bailey 1972), and may beslightly higher in high density populations (Lembeck and
Gould 1979). In exploited populations, annual adult survival is reduced; estimates of the annual proportion of surviving adults range from 49-67% (Crowe
1975b,
Fredrickson and Rice 1979, Hamilton 1982, Parker andSmith 1983, Fuller et al. 1985, Rolley 1985, Knick 1990).
The highest mortality rate- 8 1%-was found for a legally
harvested Minnesota population where illegal hunting was
believed to be high (Fuller et a2. 1985). In general, harvest mortality is greatest among the yearling and l-2 yearold young adult age classes (Govt. of U.S. 1983a, Litvaitis
et al. 1987, Knick 1990).
Longevity: (W) 12-13 years (Bailey 1974, Crowe 1975a,
Blankenship and Swank 1979); (C) up to 25-33 years(Carter 1955, Jones 1977).
Habitat and Distribution
The majority of the world’s

bobcats are found in theSpecies

range

tza

.
cl

Protected(available

area where species
for U.S.A. only)

occurs:igure 13. Distributionof the bobcat (L. rufus).
1. Mt. Agamenticus
Nature Reserve;
2. Shenandoah
II; 3. Great Smoky Mountains
II**; 4. Everglades
II#; 5. Felsenthal
IV; 6. Tishomingo
IV; 7. Aransas IV; 8. Big Bend II*; 9. Guadalupe
Mountains
II +CarlsbadCaverns
II; 10. Audubon
IV; 11. Wind Cave II; 12. Devil’s Tower V; 13. Chaco Culture V;
14. Bryce Canyon II; 15. Yellowstone
II# complex;
16. Great Sand Dunes III; 17. Kofa IV; 18. Joshua
Tree III; 19. Sequoia II complex; 20. Death Valley III; 21. Lassen Volcanic II; 22. Redwood
II**; 23. Hart
Mountain
IV; 24. Frank Church-Riverof No Return II; 25. Olympic II# (U.S.).

142

Pat? I: Species

tor in the species’ northern distribution. Marston (1942)
and McCord (1974) found that snow depths of over 15 cm
restrict bobcat movements. In areas where the two catsmet, such as Nova Scotia’s Cape Breton Island, the moreaggressive bobcat has displaced the lynx (Parker et al.
1983). A northward expansion of the bobcat’s range hastaken place over the past century, along with a corresponding northward retreat of the southern boundary of thelynx’s range, in association with the clearing for agricultureof mature conifer forests in the region (Rollings 1945,
Banfield 1974, Govt. of U.S. 1983b).

Accounts.

Chapter

5. The Americas,

Bobcat1983, Govt. of U.S. 1983a, 1992). In 1988, 37 states andfive Indian groups were authorized by the U.S. government to export bobcat pelts (Anon. 1989a). On a regional
level, in the U.S. the bobcat is totally protected in 10 states;in Canada, nowhere; and in Mexico, hunting is regulated infive states, and shooting of suspected livestock predators ispermitted on a limited basis (Govt. of U.S. 1983a, Govt. of
Canada 1983; M. Aranda in litt. 1993).
The degree to which the bobcat has been studied andmanaged in North America on both the local and national
levels makes it probably the most thoroughly examined
species in international trade today (Thomsen andLuxmoore 1990, Johnson 1990).

Population Status
Global: Category 5a. Regional: Category 4. IUCN: notlisted. In the early 1980s state wildlife authorities estimated the total U.S. bobcat population to range between
725,000 to 1 million adult animals (Govt. of U.S. 1983a).
Bobcats have historically been less abundant in the eastcentral U.S., owing to high human population density andintensive, large-scale agriculture (Peterson and Downing1952). Bobcats are also considered to be generally widespread and healthy in the Canadian (Govt. of Canada
1983) and Mexican (Govt. of U.S. 1992, M. Aranda in Zitt.
1993) parts of their range, although hunting and trapping
may have led to some local depletions (Gonzlez and Lea1
1984, G. Mowat in litt. 1993). L. rufus escuinapae (Allen
1903), the southernmost race found in Mexico, was listed
on CITES Appendix I in 1973 for reasons that are not now
clear. Craniometric studies have indicated that is is not
actually a valid subspecies (Samson 1979), and on thesegrounds it was downlisted to Appendix II in 1992. The
Mexican government has described numbers of bobcat in
the country as “adequate” (Govt. of U.S. 1983a, 1992).
Reported bobcat densities range from l-38 resident
adults per 25 km2 (summarized in McCord and Cardoza
1982, Anderson 1987; Koehler and Hornocker 1989). The
southeastern U.S. and California coastal regions appear to
support the most dense populations, probably due to highenvironmental productivity. Reduced density is associatedwith harsher environments, such as southwestern deserts
and the more northern portions of the bobcat’s range (summarized in Rolley 1987). Mean home range estimates varyfrom 0.6-326 km2 in size (summarized in Anderson 1987),
and tend to be larger in the nsrth. For example, Berg
(1979) found male ranges averaged 62 km2 in Minnesota,
whereas male ranges in Alabama averaged 2.6 km2 (Miller
and Speake 1979). Male home ranges are generally 2-3
times larger than female ranges and overlap those of several females. Male ranges overlap partially with those ofother males, while female ranges are more exclusive.

Occurrencein Protected Areas
Bobcats are likely to be found in nearly all protected areas
within their range, but confirmed records were available
only for U.S. parks and reserves (WCMC, unpubl. data).
Refugia play an important role for harvested populations,which are maintained primarily by immigration (Knick1990). Knick (1990) developed a population model
which predicted that the size of refugia needed to maintaina harvested population should be large enough to enclose
3-5 bobcat home ranges. Flores-Villela and Fernandez(1989) point out that dryland scrub habitat, important forbobcats in Mexico, is under-represented in the protectedareas system.
Principal Threats
In the last 20 years, the bobcat has been the most heavily
harvested and traded of the cat species. World demand forbobcat fur rose gradually in the late 1960s and early 1970s
and jumped in the mid- 1970s after CITES entered into
force, when the pelts of cats listed on Appendix I became
legally unobtainable for the commercial fur trade (see PartII Chapter 4). Prices offered trappers for bobcat pelts
increased sharply from a pre-1970 high of $20 to between$200-300 and as high as $600 in 1979 (Nilsson et al. 1980,
Johnson 1990). The number of bobcats killed annually in
the U.S. consequently climbed to over 90,000 in the 1980s(Govt. of U.S. 1983a); in the 1950s and 1960s about
10,000 bobcats were taken annually in the U.S., increasing
to about 44,000 in the 1970s (Johnson 1990). From 19761983, Canada reported an average annual harvest of 3,293
bobcats (Govt. of Canada 1983, Shieff and Baker 1987).
Although Mexico permits hunting, there is essentially no
documented international trade in bobcats from that country (Govt. of U.S. 1992, WCMC unpubl. data).
The financial importance of the trade to the North
American range states led to a dramatic rise in research,
particularly in the U.S., with a ten-fold increase from the
1960s to the 1970s in studies on population structure, status and distribution (Anderson 1987). Despite the volume of research, there is still concern over whether

Protection Status
CITES Appendix II. National legislation: hunting andtrade regulated throughout its range (Govt. of Canada

143

Part I: Species

Accounts.

Chapter

5. The Americas,

Pampascatcommercial trapping as practiced in North America is sustainable (see Part II, Chapter 4).
At present, trade in bobcat pelts is declining. Beginningin 1988, both harvest and export of bobcat pelts dropped
due to both market shrinkage and market saturation overseas. In addition, the European Union (formerly Community) has announced that, after 1995 (now postponed
to 1996) imports of 13 species of wild fur, including bobcat, will be prohibited unless the producer country has
either banned the use of the leghold trap or adopted
internationally accepted trapping standards (Council
Regulations EEC No. 3254/91). Europe is the primarymarket for bobcat pelts, importing 92% of North
America’s total overseas exports in 1990 (WCMC unpubl.
data). Leghold traps are the main commercial hunting
technique used to catch bobcats in North America (Baker
and Dwyer 1987, IFTF 1989).
Bobcats are generally not persecuted as pest species inNorth America. They may occasionally raid poultry, butbobcat depredations have always been uncommon,
although bounties were offered by state governments sincethe 1700s in the U.S. and throughout much of this century(Rolley 1987). In central Mexico, however, the bobcat is
reputed to be a major predator of sheep (Govt. of U.S.
1983a), and persecution by ranchers is more frequent(Woloszyn and Woloszyn 1982, Gonzlez and Lea1 1984).
The dry scrub and oak and pine forest habitats used bybobcats in Mexico have suffered the highest rates of transformation and degradation relative to other habitat types(Flores-Villela and Femandez 1989).

striped. In the Argentine pampas, the coat is longer, of amore yellow-browncolor, with muted pattern (Cabrera
and Yepes 1960). A male from central Brazil was of rusty
color with dark and conspicuous irregular black stripes
over its entire body when young (three months old), but by
the age of eight months, stripes were visible only on thelimbs and underparts (Silveira in submission).
A taxonomic evaluation of 96 museum specimens leads
Garcia-Perea (1994) to propose that, given pronounced
geographic differences, the “pampas cat” is actually three
species: Lynchailurus pajeros (Desmarest, 18 16) [highAndes from Ecuador to Patagonia and throughout
Argentina]; L. braccatus (Cope, 1889) [warm grasslandand sub-tropical forest in Brazil, Paraguay and Uruguay];
and L. colocolo (Molina, 1782) [central and northwestern
Chile]. She plans to test the degree of differentiation
between the three types by molecular analysis.
The long hairs on the pampas cat’s back (up to seven
cm in length) form a sort of dorsal mane: when it sets its
hairs on end, it looks larger than it really is, which is only
a little bigger than a domestic cat. The recorded wildcaught weights range from 3-3.7 kg (n=3: Redford andEisenberg 1992, Silveira in submission). In captivity, they
have weighed up to 7 kg (Green 1991).
Although the pampas cat is relatively common andwidespread, there is surprisingly little data available on its
ecology. It has been described as preying chiefly on small
mammals such as guinea pigs (Cabrera and Yepes 1960,
Guggisberg 1975, Ferrari et al. 1984, Rabinovich et al.
1987, D. Brooks in litt. 1989), as well as ground-dwellingbirds-forexample, in Patagonia pampas cats have been
observed to take penguin eggs and chicks from nests (D.
Boersma in Zitt. 1990). Pampas cats are thought to be predominantly nocturnal and terrestrial. However, they havebeen observed in daylight hours in the wild, and a malepampas cat kept in Brazil’s Parque Zoologico de Goiania
showed great skill in tree climbing, and spent most of its
resting periods draped over the highest fork of a small deadtree installed in its cage (Silveira in submission).

Action Planning
Projects 17 and 104.

Pampas cat, Oncifelis
co/oco/o (Molina, 1782)
Other Names
Chat des pampas (French); Pampaskatze (German); gatopajero, gato de 10s pajonales, osio (Spanish); gato depajonal (Argentina, Ecuador, Peru, Paraguay); gato montes (Bolivia, Chile, Ecuador, Peru, Paraguay); gato peludo(Bolivia); gato palheiro (Brazil); gato colocolo (Chile);
gatillo (Ecuador); osjollo, chinchay (Peru).

Biology
Reproductive season: (C) April-Julysphere) (Green 199 1).

(northern

hemi-

Litter size: (C) 1.3 1 t 0.13 (n=13) (Mellen 1989); range
l-3 (Rabinovich et al. 1987).
Age at first reproduction:
Eaton 1984).

Description
and Behavior (Plate 4)
The pampas cat has a wide distribution and broad habitat
selectivity, and its appearance varies in different parts of its
range. In the high Andes, it is grey-colored with reddish
stripes broken up into spots, and looks rather similar to
the Andean mountain cat, although it is not so heavily(C) two years

n= 1 female:

Longevity: (C) average nine, but up to 16.5 years (n=3)(Prator et al. 1988).
Habitat and Distribution
The pampas cat is strongly associated with grass and shrub

144

Par? I: Species

Accounts.

Chapter

5. The Americas,

Pampascat

to over 5,000 m, where it is possibly sympatric with the
Andean mountain cat (Grimwood
1969, Redford andEisenberg 1992).
The range portrayed in Fig. 14 assumes that pampas cat
populations are largely continuous. However, in keepingwith her theory that there are actually three different speciesof pampas cat, Garcia-Perea (1994) has produced an alternative range map showing disjunct distributions (Fig. 15).
The range she portrays in south-central Brazil should be
extended to the east as shown in Fig. 14, based on specimen
records collected by Silveira (in submission).
Population Status
Global: Category 5a. Regional: Category 4. IUCN:
Indeterminate. The pampas cat is widely distributed, tolerant of altered habitat (including secondary growth, forest
plantation, and the fringes of agricultural and settled areas:
P. Crawshaw, C. Weber in litt. 1993), and international
trade in its pelt ceased in 1987 (WCMC unpubl. data). In
the Paraguayan Chaco, it has been described as less common than the Geoffroy’s cat (Brooks 1992). Although

Protected area where
species may occur

Figure 14. Distributionof the pampas
cat (0. co/oco/o).
1. Sangay II** (Ecuador);
2. Huascarn
II#; 3. Manu II# (Peru);
4. Amboro
II complex;
5. Eduardo Avoroa IV (Bolivia);
6. Defensoresdel Chaco II (Paraguay);
7. Serra da Capivara;
8. Aparadosda Serra II; 9. Emas II (Brazil); IO. Dunas de Cabo
Polonio Ill (Uruguay);
11. Lauca II* complex;
12. Bosque Fray
Jorge II*; 13. Torres del Paine II* complex(Chile); 14. Los Andes
I; 15. La Florida V complex;
16. Lihuel Calel II complex(Argentina).

chailurus

habitats (Fig. 14). In addition to the pampas grassland
formations for which it is named, it also occurs throughout the cerrado (open wood, shrub, and grass complexes)of central Brazil (L. Silveira in submission). The pampas
cat also occurs in several forest types, typically open
woodland or scrub thicket, such as the Gran Chaco, butalso the belt of “yungas” cloud forest that runs along theeastern slope of the Andes (Cabrera 196 1, Grimwood
1969, Cabrera and Willink 1980). It is absent only fromlowland rain forest, both tropical (Emmons 1990) and temperate Valdivian (Taber et al. 1974). At the southern
extent of its range, it occurs in the cold semi-arid desert ofPatagonia. In Uruguay, it is found in low-lying swampy
areas with clumps of long esparto grass (Ximenez 1961),
and also occurs around the Pantanal floodplain (L. Silveirain submission). It occurs on both the eastern and western
slopes of the Andes, with an elevational range from 100 up

Figure 15. Distributionof proposed
three
cat according
to Garcia-Perea(1994).

145

speciesbraccatusof pampas

Part I: Species

Accounts.

Chapter

5. The Americas,

Jaguarundi

unpatterned fur. The jaguarundi also differs from the othersmall cats of Latin America by its elongated rather thanrounded head (Eisenberg 1990). Reported adult weights
range from 2-9 kg (Mondolfi 1986, Guggisberg 1975). In
Belize, two males averaged 5.9 kg and two females 4.4
kg (Konecny 1989). There are three different color forms,
which may sometimes occur in the same area or even thesame litter (Konecny 1989, Brooks 1992)-black,brownish grey, and red. In general, however, the darker colors
are most commonly associated with inhabitants of rain forest habitats, while the paler color is found most frequently
in drier environments (Emmons 1990). The red form wasonce considered a separate species F. eyra (Fischer, 18 14).
It has been suggested that the jaguarundi prefers to hunt
ground-dwellingbirds rather than mammals (Gaumer
1917, Leopold 1959, Hall and Dalquest 1963), and analysis of 23 stomachs from Venezuela (Mondolfi 1986, Bisbal
1986) shows that birds are frequently caught (found in 5470% of the stomachs). Rodents, rabbits, and reptiles werealso found in 40-51% of the stomachs. In Belize, scat
analysis indicated that arthropods are frequently eaten(remains found in 72% of scats); birds occurred in 22% ofscats and rodents in 95% (Konecny 1989). Jaguarundis
have also been observed to prey upon characid fishstranded in a puddle (Manzani and Monteiro 1989).
Rengger (1830) suggested that the solid coat of thejaguarundi is probably associated with the fact that thesecats hunt more diurnally or terrestrially than spotted felids.
While nocturnal activity (as well as arboreal foraging) is
occasionally observed (Leopold 1959, Guggisberg 1975,
McCarthy 1992), Konecny’s (1989) radiotelemetry studyof four jaguarundis in Belize found the period of peak
activity to be 0400- 1100, with only residual activity(movements of less than 100 m/hr) after sunset.
Jaguarundis have been frequently observed travelling or
foraging in pairs (Rengger 1830, Guggisberg 1975,
McCarthy 1992).

pronounced extinct in Uruguay over 30 years ago (Cabrera
and Yepes 1960), it probably still exists, but very sparsely
due to draining of wetlands for ranching and agriculture(A. Ximenez in litt. 1991). Although it appears to have awide range in Brazil (the cerrado is Brazil’s second largest
habitat type after tropical rain forest: Olson et al. 1983),
records are scarce and the species is generally consideredrare (L. Silveira in submission). The pampas cats of Chile(L. coZocolo of Garcia-Perea [ 19941) are the most endangered group due to small geographic range.
Protection
StatusCITES Appendix II. National legislation: protected acrossmost of its range. Hunting prohibited: Argentina, Bolivia,
Chile, Paraguay. Hunting regulated: Peru. No legal protection: Brazil, Ecuador. No information: Uruguay (Fuller
et al. 1987).
Principal Threats
The pampas region of Argentina and Uruguay has beenheavily settled and grazed relative to other regions, and thestatus of the species should be investigated here. The pampas cats of Argentina were formerly hunted in large numbers for the fur trade-78,000
skins were exported from
1976- 1979 (Mares and Ojeda 1984)-but
international
trade has ceased following a last shipment of 10,000 pelts
exported in 1987 to clear old stocks (WCMC unpubl.
data). L. Silveira (in submission) reports that the pampas
cat is known to raid chicken houses occasionally.
Action Planning
Projects 99 and 105.

Jaguarundi,
yaguarondi

Herpailurus(La&p&de, 1809)

Biology
Estrus: (C) 3.17 -+ 0.75 days (n=6).

Other Names
Otter cat (English); jaguarondi (French); Jaguarundi,
Wieselkatze, Eyra (German); yaguarundi, onza, gatomoro, gato eyra (Spanish); halari (Belize); maracaja-preto,
gato-preto, gato mourisco (Brazil, Uruguay); gato griz
(Bolivia); gato pardo, gato servante, ulama (Colombia);leon brefiero (Costa Rica, Peru); jaguarondi, chat noir
(French Guiana); tejon, mbaracaya-eira (Guatemala); gato
cerban (Honduras);
kakicoohish(Kekchi);ekmuch(Mayan); tigrillo congo, tigrillo negro (Panama); leoncillo,anushi-puma (Peru); boesikati (Suriname); gato Cervantes(Venezuela).

Estrus cycle: (C) 53.63 t 2.41 days (n=8: Mellen 1989).
Gestation: 70-75 days (Hulley 1976, P. Andrews
1993).

in litt.

Litter size: (C) 1.83 t, 0.24 (n=12: Mellen 1989); range l4 (Hulley 1976).
Age at sexual maturity:
Andrews in Zitt. 1993).

(C) 2-3 years (Hulley

1976, P.

Longevity: (C) up to 15 years (Prator et al. 1988).
Description
and Behavior (Plate 4)
The jaguarundi has a distinctly weasel-like appearance,with its elongated slender body, short legs, and sleek

Habitat and Distribution
A cat of the lowlands not generally found above 2,000 m

146Part I: Species(Vaughan 1983), the jaguarundi otherwise occupies abroad range of both open and closed habitats-fromdry
scrub, swamp, and Savannah woodland to primary forest.
In Venezuela, it has been most frequently collected in tropical dry forest, relative to other habitat types (Bisbal 1989).
Jaguarundis are more rare and thinly distributed in moist
forest types, especially deep rain forest (Konecny 1989,

Accounts.

Chapter

5. The Americas,

Jaguarundi

L. Emmons in Zitt. 1993). Jaguarundis have been reported
to prefer forest edges and secondary brush communities(Bourliere 1955, Mondolfi 1986), but this may be because
it is in such areas that these primarily diurnal cats are mostfrequently seen. In Belize’s Cockscomb Basin Wildlife
Sanctuary, Konecny (1989) found that jaguarundis aremost frequently associated with riparian and old field habi-

Figure 16. Distributionof the jaguarundi(H. yaguaronc0).
1. Laguna Atascosa
IV (Texas, U.S.); 2. Cockscomb
Basin IV (Belize); 3. Volcn de Pacaya Ill(Guatemala);
4. Rio Platano II* (Honduras);
5. Braulio Carrillo II complex(Costa Rica); 6. Barro
Colorado
II (Panama);
7. lsla de Salamanca
II (Colombia);
8. Guatopo
II (Venezuela);
9. Purac* II(Colombia);
10. Pacaya-Samiria
VIII; 11. Manu IN (Peru); 12. Manuripi
Heath IV complex (Bolivia);
13. lguazu II** (Argentina)+ Iguacu II** (Brazil) complex;
14. Lihuel Calel II complex (Argentina).

147

Part I: Species

Accounts.

Chapter

5. The Americas,

Jaguarundient, widely spaced portions of their ranges for irregularperiods of time, rather than making regular boundary
patrols (Konecny 1989).

tats. Access to dense ground vegetation appears to determine habitat suitability for the jaguarundi, but of all thesmall New World felids, it is most flexible in its ability tooccupy diverse environments (Fig. 16).

Protection
StatusCITES Appendix II; Central and North American populations Appendix I since 1987. National legislation: protected over much of its range. Hunting prohibited:
Argentina, Belize, Bolivia, Colombia, Costa Rica, French
Guiana, Guatemala, Honduras, Mexico, Panama, Paraguay,
Suriname, Uruguay, United States, Venezuela. Hunting
regulated: Peru. No legal protection: Brazil, Nicaragua,
Ecuador, El Salvador, Guyana (Fuller et al. 1987).

Population Status
Global: Category 5c. Regional: Category 5. IUCN: notlisted. The jaguarundi appears to be relatively common
over much of its range (Koford 1976, Vaughan in press)
although, while present throughout the Amazon basin rain
forest, it is more rare in this habitat type (L. Emmons ipzZ&.
1993). The jaguarundi may now be extinct in Uruguay(Thornback and Jenkins 1982), and is very rare in thesouthwestern United States near the Rio Grande (Tewes
and Everett 1986).
In Belizean rain forest, home ranges for malejaguarundis were very large (Konecny 1989), several times
larger than those reported for sympatric jaguars weighingnearly 10 times more (Rabinowitz and Nottingham 1986).
One female used a home range that varied between 13-20
km2, while two males used home ranges of 100 and 88
km? The home ranges of the two males overlapped lessthan 25%. Both sexes exhibited a pattern of using differ-

Principal Threats
Generally not exploited for commercial trade, althoughjaguarundis are doubtless caught in traps set for commercially valuable species and may be subject to low intensity hunting pressure around settled areas. They arenotorious for predation on domestic poultry (Rengger
1830, Alvarez de1 Toro 1952, Leopold 1959, Hall and
Dalquest 1963, Goodwyn 1970, Koford 1976, Ferrari et al.
1984, Bisbal 1986, McCarthy 1992).

148Part II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

Part II
Major Issues in Cat Conservation
Chapter 1
Cats and Habitat Loss
Introduction

Habitats for CatsThis planet’s land area amounts to over 148 million km2.
With the total human population at over 5.3 billion, thisamounts to a density of more than 40 people per km2(excluding 18 million km2 of the Antarctic land mass andthe area taken up by lakes). The human population has
more than doubled since 1950, and is projected to reach
8.5 billion by 2025. Despite a worldwide decline in thebirth rate since 1970, demographers say that the total population will continue to increase simply because more people than ever before will reproduce. It is projected tocontinue growing over the next 200 years, eventually stabilizing at 11.6 billion in 2200-or about 90 people per
km2 (WRIKJNEPKJNDP
1992).
In order to support this growth, wild lands are coming
under increasing use and development. Figure 1 shows thedegree of human modification around the globe. Loss andfragmentation of habitat are often the primary cause of
species extinctions and the decline of biodiversity(WCMC 1992). The increase in the number of people in
the world and the decline in the amount of wilderness area
will continue unless fundamental changes are made in our
economic and social systems globally, nationally, andlocally, to make the present way of development into asustainable process (IUCN/WWF/UNEP
199 1).
This chapterhastwo primary purposes. The first is topresentan overview of the statusof the world’s habitats,and to evaluate the implications for cat conservation.
Critical habitat types for cats are identified, and global
trends in habitat loss and fragmentation are reviewed.
Given the socio-economic pressuresreferred to above,
thereis a certainamountof inevitability in thesetrends,butthere is alsoroom for optimismregardingthe potential toconserve cats in a variety of human-modified habitats.
This will require that cat specialiststake an active role indevelopment-relatedwork which lies outsidethe realm of
traditional researchactivities. The secondpurposeof thischapter is to review the protected areasof the world intermsof cat conservation,and focus on what changesare
neededto makethem moresuitablehavensfor cats.

A global overview of the world’s habitats in relation tocat distribution calls for habitatsto be grouped according
to similarities. There is currently no universally recognized global habitat classification scheme. The various
schemesare basedon either (1) distinct communities ofplant species(taxonomy) or (2) similaritiesin plant physiognomy (structure: forest, grassland),phenology (evergreen, deciduous), or environmental variables (altitude,
climate, hydrology) (WCMC 1992,Anon. 1993d). One of
the latter types of classificationsis usedhere becauseit isclear, generally speaking,that the physical nature of theenvironment hasbeen a powerful selective force shaping
cats (Eisenberg 1981: 211). Examples include the snow
leopard’spale coat, the arborealmargay’sflexible ankles,
and the sandcat’s fur-covered paws. Cats have a rather
indirect relationshipwith plant species,unlike their mainly
herbivorous prey. It is expected that when the world’s
scientists finally agree upon and develop a universally
acceptedhabitat classification scheme,vegetation structure will be the primary basis(Anon. 1993d). It is hopedthat this sort of classification adequatelyreflects the way
catsperceive and selecthabitat.
The map of major world ecosystemcomplexesdeveloped by Olson et al. (1983) (Figs. 2a-fj usesa classification system developed principally to refine global
estimatesof the amountof carbon containedin plants, sothat similaritiesin vegetative structure (rather than species
composition) are emphasizedin the definition of habitat
types. Unlike other major habitat classifications, large
areasof human-modifiedenvironments,suchasrice paddiesor settledareas,are includedasdistinct “ecosystems.”
This is of key importance, ashuman-modifiedvegetation
comprisesmost of the world’s land surface,far out-stripping natural, original vegetation (Fig. 1; Anon. 1993d:9).
The definition of each habitat type is given in Table 1,
and Figs. 2a-f map the global distribution of thesehabitat
types asof 1980. Table 2 lists habitat types, their globalarea, and the number of cat speciesassociatedwith each
type. As a componentof cat speciesvulnerability ranking,149

Part II: Major

issues.

Chapter

1. Cats and Habitat

Loss

Box 1
Habitat Classif ication and Species Associations
Olson ef al. (1983) originally designated 47 land-based
habitat types, which have been combined here to yield atotal of 21. For example, where their original map distinguished three sub-types of temperate broad-leaved
forest, they are combined here. Type definitions are
given in Table 1. Each cell or pixel in the map represents an area of 0.5 degrees latitude by 0.5 degrees longitude, or 55 x 55 km at the equator = 3,025 km*. Usinga variety of vegetation and land-use maps and data
sources, Olson et al. (1983) assigned each pixel to its
predominating
habitat type. To reduce bias, several
habitat types were defined as vegetative mosaics, suchas 2c, other dry woods/scrub/grasscomplexes.
Withsuch a large pixel size, however, loss of detail is
inevitable. As discussed later in this chapter, most of theworld’s protected areas are well under 3,000 km* in size.
If surroundedby human-modified
landscape,thesepatches of natural habitat generally do not show up on
the map.

tion was assigned according to relative abundance ofa species within a given habitat type. Degree of association was also based upon how large a proportion of a
species’ total contemporary range is taken up by a partitular habitat type. Relative abundance
and relative
area are not always congruent.
For example, northern
taiga forest (habitat type Zd) occupies more of theEurasian lynx’s range than do southern taiga and otherconiferous forest (habitat types 1 a and 1 b), but the lynx
occurs at greater densities in the latter (Matjuschkin1992). Similarly, although broad-leaved
humid forest(habitat type le) occupies a major portion of thejaguarundi’s range, the jaguarundi is sparsely distributed
in dense primary forest compared to more open habitats (L. Emmons in liiy. 1993). There were not sufficient
data available to use either relative abundance or area
consistently.
However, to compensate as far as possible, assignment of degree of association in this study isconservative.

Species-habitat
association
was defined as either
strong, significant, marginal or absent. Habitat associations are given for each species in Appendix 4. Whensufficient reference material was available (references
are given in the Species Accounts), degree of associa-

In addition, a habitat type which occupies only a small
portion of a species’ range was defined as significant
if it harbors an important sub-population:for example,
coniferous forest (habitat type 1 b) for the Siberian tiger,or mixed woods (1 d) for the north African serval.

trees: black-footed cat, sandcat, Andean mountain cat,and manul.
Mammalian speciesrichness is greatest in tropicalmoist forest (Eisenberg X981), which covers only about
7% of the Earth’s land area,but constitutes a major partof the rangesof nearly half of the cat species.Both tropical humid forest (habitat type le) andgrasslandand shrubland (habitat type 4a) each have 14 species strongly
associatedwith them-almost twice as many asthe next
most species-rich habitat type, tropical Savannah andwoodland (2a), with eight speciesstrongly associated.
Comparedto other habitat types, however, tropical humid
forest is notable for its lack of specieswhich are marginally or significantly associated. Only three speciesare
significantly associatedwith tropical humid forest-thejaguarundi,jungle cat, and oncilla. The jaguarundi occurs
throughout the humid forestsof Latin America, but at low
densitiesin denseprimary forest, while the oncilla appears
to be absent from the lowland humid forests of the
Amazon basin, although it is presentin humid forests tothe north. The jungle cat is alsothinly distributed in drier

species-habitatassociationswere defined as strong, significant, marginal, or absent(seeBox).
Generally speaking, cats make use of a variety ofbroadly defined habitats. The averagenumberof habitats
in which cat speciesoccur is 7.5 (SD=3.49; mode=7;range= 2- 15), or just over one-third of the 21 habitat types. The
averagenumberof strong and significant habitat associations per speciesis 4.6 (SD=2.I9; modes=2& 4; range =1- lo), or about one-fourth of the total available habitat
types. Only one habitat type, polar ice and desert (5b),lacksa memberof the cat family. L
From the datain Table 2, the importanceof forest habitat to catsis immediately evident. Closedandopen woodlandscover approximately 40% of the Earth’s land area,and harbor 89% of the cat species (32 of 36 species).
Closedforest is particularly important habitat. This makes
sense:cats, with their adaptationsfor climbing and stalking, are believed to have evolved from a forest-dwelling
miacid ancestor(Ewer 1973, Leyhausen 1979,Eisenberg
1986). Only four speciesare adapted to live wholly insparselyvegetatedenvironmentscharacterizedby lack of150

Pat-t II: Major

1. Humandisturbanceto naturalvegetation.

Source:Chapter

1. Cats and Habitat

Lossropical montane complexes (2b), and high alpine tundra
5a). These three habitats share the features of being smalln overall area, and locally patchy or insular. With the
exception of high alpine tundra, all have decreased in area
over the 198Os, as discussed below.

parts of the humid forests of tropical Asia. Still, tropicalhumid forest makes up large portions of these species
ranges-no cats are marginally associated with this habitat
type. In general, it seems that cats are either strongly tied
to rain forest, or otherwise not at all.
While the overall felid trend is toward broad habitat
selectivity, a substantial minority are more specialized. As
shown in Table 3, 14 species (39% of the family) are classified as narrow in habitat association, being associatedwith a total of six or fewer habitat types. This point should
be qualified, however, by noting that the majority of theseare small, poorly studied cats (e.g., compare with Table 1
on research effort in Chapter 3), and future field work isneeded to confirm habitat associations.
Of these 14
species, seven, or 50%, are strongly or significantly associated with tropical rain forest, a proportion very similar
to that for the cat family as a whole (47%). In other words,
habitat specialists are not more likely to be strongly associated with tropical rain forest than more generalist cats.
Habitat specialization does not necessarily correlatewith vulnerability, however. Considering the 19 species
ranked in Categories 1-3, which are of highest global conservation priority, it can be seen that a slightly greater proportion, when compared with all felids, is associated withtropical rain forests. Table 4 lists habitat types with thehighest percentages of Category l-3 cats closely (strongly
and significantly) associated. In addition to tropical moist
forest, other habitats which are relatively more importantfor vulnerable cats include major wetlands (habitat type 6),

FigureIssues.

Habitat Loss andFragmentation:
An
Overview of Global Trends
Habitat Loss
There are currently no global systems which monitor the
status of a standardized, globally recognized set of habitat
types, although the conservation community has recognized the need for such a system and made its developmenta priority (Anon. 1993d). Detailed quantitative data isavailable, in reality, only for tropical forests (C. Billington,
World Conservation Monitoring Centre, pers. comm.
1994), and the “state of the art” is the United Nation’s
Food and Agriculture Organization’s Forest Resources
Assessment 1990 Project (FAO 1993; J. Blockhus, IUCN
Forest Conservation Programme, pers. comm. 1994).
Tropical moist (includes rain forest) and dry forests harbor 68% of the vulnerable Category l-3 cats (13 species),and 64% of all the species in the family Felidae (23species; see Table 7). Moreover, as pointed out above, cats

Conservation

151

International.

Part II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

Table 1
Key to Global Habitat Types
Modifiedfrom Olsonet al. 1983

RegionNumberVegetation Type

All

1.

Closed460

Taiga and other conifer

ATE

1 (a)

Main and southern taiga. Coniferous evergreen trees of the boreal climate zone,
mixed toward the southern boundary with deciduous birch, poplars, and larch.

AE,O

1 (b)

Other conifer. Mostly evergreen needle-leaved trees, excluding
both regions of persistent snow and warm to hot climes.

A,E,N, W

Mid-latitude

NvJ,o

1 Cc)

Temperate

A,uLT

1 Cd)

Mixed woods. Deciduous to evergreen
planted conifers interspersed.

A, TW

Tropical/subtropicalbroad-leaved

A,-LSO1 (e)Broad-leaved humid forest. Includes both evergreen
deciduous semi-evergreen(monsoon) forest.

A,TW

l(f)

Tropical dry forest and woodland. Drought-deciduous
forest with many dry
months alternating with rainy season (or two near equator). Drier or mostburned parts often have grassy field layer, transitional to savannahs.

All

2.

Open or Interrupted

w-s,0

aa)

Tropical Savannah and woodland. Palms and other trees and shrubs scatteredin grassy undercover, frequently burned or grazed.

A7-w

2(b)

Tropical montane complexes. Rainy or cloud forest situated at I ,OOO-2,500
meters (or higher) with alpine meadow, scrub, rock interruptions; also highlandcrop, burn, and fallow areas.

Allbroad-leaved

Forest and Woodland(Figure 2a)most taiga, inand mixed forest
broad-leaved

forest. Deciduous

hardwoods,

broad-leaved

localized

or no conifers.

forest, with native orforestWoodlandrain forest and seasonally(Figure 2b)

Other dry woods/scrub/grasscomplexes. A broad category including
Mediterranean-type
sclerophyllic vegetation, succulent and thorn woods andscrub, other dry or highland tree or shrub types (juniper, open pine) and quebracho forest of northern Argentina and Paraguay.

ATE

2(d)

Northern or maritime taiga. Mostly stunted and patchily distributed
conifer, sub-alpine.

All

2(e)

Second growth woods and field mosaics. Includes more than 40% closed forest
or open woods, mostly heavily disturbed by cutting or fire, or regrowing or
replanted, alternating with trees or other crops, settlements, cultivation or open
areas (mainly temperate, 72% of total area). Also includes predominantly cultivated or grazed landscapes with isolated remanants of forest or more open
woodland (mainly tropical, 28% of total area).
Continued

152

borealon next pagePart II: Major

Issues.

Chapter

RegionNumberVegetation

AU

3.

Settled Areas: Cropped, Residential, Commercial, andAssociated Marginal lands (Figure 2~)

-LE

3(a)

Paddyland. Rice and tropical crops, with settlementsinterspersed.

I. Cats and HabitatType

and/or tree remnants

Towns, farms, and other irrigated dryiand row crops.

Ail

All

4.

Grass and Shrub Complexes: tow Vegetation with Few
or No Trees (Figure 2d)

Ail

4(a)

Grassland or shrubland. includes most types of shrub and grassland (prairie,
pampas, steppe, puna: 81% of total area), aipine meadow (18%), and heathand mooriand (1%).

E

4(b)

Cold grass or stunted woody complex. includes Tibetan and Siberian meadows(33% of total area), and dwarf and scrub vegetation at polar and aititudinai limitsof taiga or montane forest or woodland (67%).

w,ww
ww

5.

Tundra and Desert (Figure 2e)

E,A,J’J

W)

Very cold areas with sparsevegetation
Tundra. Dwarf scrub, tussock, sedge and other herb patches, commonly overpermafrost. includes both classical Arctic tundra as well as sparsely vegetatedhigh alpine areas.
Polar desert. Bare rocks, lichens, glacier and permanent
Arctic and Antarctica.

&Asnow fields of high

W,S,W

Desert and semi-deserf

W

5(c)

Cool semi-desert scrub. Tail or low shrub, semi-shrub, shrub steppe andopen grassland in comparatively cool, continental climates.

NW,0

5(d)

Sand desert. Bare, moving sand dunes, with sparse shrub or grass cover.

J’LEW

w

Other desert and semi-desert. Mostly warm to hot arid shrublandwith some succulents and ephemerals.

All

6.

,

Major Wetlands (Figure 2f)
includes swamp/marsh,of cool or cold climates.

A = The Americas
E = Eurasia
N = North Africa/Southwest
0 = Oceania
S = Sub-SaharanAfricaT = Tropical Asiaand grassland,

Asia

153

mangrove

and tropical swamp woods, and bog/mire

LossPart II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

Table 2Cat Species Occurrence in Major World Habitat Types
Habitat Type’Global Area(106 km*)%Number of Species Associated*
Str Sig Mar Tot [% ofFamily]Closed and Open Forest
and Woodland (1 & 2)

58.17

[39.3%]

29

1

2

32

[89%]

Closed Forest and Woodland (1)30.79[20.8%]

19

8

5

32

[89%]

Mostly taiga and other conifer (I a & lb)

10.66

[72%]

3

4

5

12

[33%]

Main and southern taiga (1 a)7.16[ 4.8%]

2

0

I

3

[ 8%]

Other conifer (1 b)3.50[ 2.4%]

3

4

5

12

[33%]

503

[ 3.4%]

3

9

8

20

[56%]

1.49

[ LO%]

0

6

6

12

[33%]

3.54

[ 2.4%]

3

7

9

19

[53%]

15. IO[10.2%]

17

5

I

23

[64%]

10.38

[ 7.0%]

14

3

0

17

[47%]

4.72

[ 3.2%]

7

9

4

20

[56%]

27.38

[18.5%]

14

16

0

30

[83%]

7.32

[ 5.0%]

8

5

5

18

[50%]

0.60

[ 0.04%]

1

6

IO

17

[47%]

7.60

[ 5.1%]

6

IO

2

18

[50%]

Northern or maritime taiga (2d)4.35

[ 2.9%]

0

2

0

2

[ 6%]

Second growth woods and field mosaics (2e)7.20

[ 4.9%]

2

15

11

28

[78%]

Settled Areas (3)15.90[10.7%]

0

6

11

17

[47%]

Paddyland

2.00[ I .4%]

0

4

1

5

[14%]

13.90

[ 9.4%]

0

4

12

16

[44%]

Grass and Shrub Complexes (4)23.90[16.1%]

14

4

II

29

[81%]

Grassland

21.40[14.5%]

14

4

IO

28

[78%]

2.55

[ I .7%]

1

0

4

5

[14%]

Mid-lat. broad-leaved
Temperate

and mixed (Ic & Id)

broad-leaved

forest (1 c)

Mixed woods (1 d)Tropical broad-leaved
Broad-leaved

forest (le & If)*

humid forest (1 e)**Tropical dry forest and woodland(If)

Open or Interrupted Woods (2)Tropical

Savannah and woodland

Tropicalmontanecomplexes

Other dry woods/scrub/grass

(2a)

(2b)
complexes

(2~)

(3a)

Towns, farms and other croplandor shrubland

(3b)

(4a)

Cold grass or stunted woody complex(4b)

Continued on next page154

Part II: Major

Global Area(106 km*)%

Habitat Type1

Tundra and Desert (5)Issues.

Chapter

1. Cats and Habitat

Number of Species Associated*
Str Sig Mar Tot p43of
Family]44.40[30.0%]

9

6

3

18

[50%]

26.20

[17.7%1

2

I

6

9

25%]

Arctic tundra and barren alpine (5a)11 .oo[ 7.4%]

2

1

6

9

[25%]

Polar ice and desert (5b)15.20

[10.3%]

0

0

0

0

-

18.20

[12.3%]

7

5

4

16

[44%]

2.00

[ 1.4%]

3

1

4

8

[22%]

5.20

[ 3.5%]

1

3

4

8

[22%]

11 .oo[ 7.4%]

4

5

4

13

[36%]

2.90

[ 2.0%]

6

1

7

14

[39%]

Very cold sparse vegetation

Desert and semi-deserf
Cool semi-desert(5a & 5b)(5c, d & e)scrub (5~)

Sand desert (5d)Other desert and semi-desert(5e)Major Wetlands (6)Key:
1 See Table 1 for definition of habitat types.
2 Str=Strongly associated. Sig=Significantly
associated. Mar=Marginallyassociated. Tot=Total. [%I=% of 36
species in the family Felidae. See Box 1 for explanation of how species-habitat
associations were assigned.

* FAO (1993) estimates tropical and subtropicalforest cover as 19.1 million km2 in 1980, a refinement of its earlier
estimate of 19.35 million km? This total differs noticeably from the 15.10 million km2 given above for habitat types
1 e & 1f, although FAO data on forest cover was a major source of information for Olson et al. (1983). However,on pp. 48-49, Olson et a/. (1983) explain that total forest cover can be calculated from their data in a way closer to
FAO’s methodology.
From habitat type 2e, Second growth woods & field mosaics, 1.7 million km2 can be allocated to 1 e, broad-leaved humid forest, and 0.6 million km2 to If, tropical dry forest & woodland.
The 0.6 million
km2 of 2b tropical montane complexes can also be added to le. [Note: the total area covered by this habitat typeis actually 1.2 million km? but Olson et al. apportion half, consisting mainly of non-woods components, to 3, settled areas.] In this way, a total coverage of 19.4 million km2 is obtained for tropical and subtropical forest, a total which agrees closely with the FAO estimate.

** Although the following sub-categories
are not mapped, Olson et a/. (1983: p. 48) provide details on the types ofhumid broad-leaved forest which make up habitat type le.

Habitat type 1e sub-category

Area (106km*)

Lowland rain forest
Mangroves (of forest structure)
Evergreen equatorial forest

3.0
0.2
2.8

Tropical seasonal forest
Evergreen or deciduous “moist” forest, closed or regenerating
Planted, degraded, poor site, or marginal “forest”
Total tropical/subtropicalbroad-leaved

humid forest

well

7.4
6.0
1.4
10.4

155

Loss

Part II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

156

Part II: Major

-j

z
7

157

&-a
i

----

Issues.

Chapter

1. Cats and Habitat

Loss

Part II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

E
E
8

158

.-k

Part II: Major

Issues.

Chapter

1. Cats and Habitat

LossVery cold areas with sparsevegetation

El Tundra

Deserta8-.’ Cool and semi-desert

Polar desert

2e. Habitat

map:

scrub

Sand desert
B Other

Figureand semi-desert

Tundraand Desert.

Source:World

Conservation

desertMonitoringand semi-desert
Centre.

Part II: Major

I

#

8

161

Issues.

Chapter

1. Cats and Habitat

loss

Part II: Major

A lioness

Issues.

among

Chaptertree stumps

1. Cats and Habitat

which

were

Loss

part of the Gir Forest,

the last refuge

which are associated with tropical forest tend to bestrongly associated (this tendency is much stronger for rain
forest than tropical dry forest), so that tropical forest constitutes major portions of these species’ ranges. Therefore,
it is worth examining loss of tropical forest habitat indetail.
Tables 5 and 6 present the FAO’s data on tropicaldeforestation by ecological zone and geographical region
over the 1980s. From Table 5, it may come as a surprise,given all the attention paid to loss of rain forest in the
media, that this habitat type is undergoing a lower rate of
deforestation than moist deciduous, tropical dry, and montane forest. Moreover, it is the tropical forest type withthe largest global area, and the type which still covers mostof its original land area (76%). (The figures for the othertypes are somewhat deceptive. In the rain forest ecologicalzone close to 100% was originally forested, while in thevery dry zone considerably less than 100% is likely to havebeen forested originally: FAO 1993). This is not to underplay the importance of tropical rain forest-the
habitattype associated with the largest number of vulnerable cat
species-but to highlight the significance of deforestationin other tropical zones. Loss of montane forest is especially significant for the oncilla in Latin America, and forthe tropical Asian cats of the Himalayan region.
Table 6 shows the regions which have undergone thehighest rates of deforestation over the past decade are continental southeast Asia, insular southeast Asia, Central
America and Mexico, and west Africa. Insular southeast
Asia still has over 100 million ha ( 1 million km”) of for-of the Asiaticlion.

est cover. The remaining three regions have between 4070 million ha remaining, asdo other regions with lower
rates of deforestation: west and eastSahelianAfrica, andsouth Asia. Tropical South America is the region withthe most remaining tropical forest cover, although comparisonwith its total land areastill indicatesthat a significant portion has already been lost. Central Africa
underwent the lowest rate (5.3%) of deforestation in the
1980s and still has 200 million forested ha, althoughnearly half of its original forest areahasbeenlost.
Table 7 lists the vulnerable cat speciesfound in theseregionswhich are strongly or significantly associatedwith
tropical moist or dry forest. It is readily apparentthat continental and insular southeastAsia, which are both undergoing very high rates of deforestation, have the largest
numberof speciesassociated,and are of top global priority for habitat conservation efforts asfar as cats are concerned. While cultivation and settlement play animportant role, much of tropical Asia’s deforestation is
the result of logging. Since the 1960s the Asia/Pacificregion hasfar outstrippedAfrica and Latin America in its
averageannualproduction of raw tropical roundwood. For
example, in 1990, production was close to 120 million
cubic meters, while Africa and Latin America each producedonly around20 million cubic meters(FAO 1993).
What is happeningto all the other types of habitat withwhich cats are associated?There is little publisheddata.
Fortunately, it is possibleto undertakea preliminary evaluation with regard to the habitat types of Olson et aZ.
(1983), the foundation of the cat speciesvulnerability

162

Part II: Major

lberian lynx, L. pardnus
Bornean bay cat, C. badia
Chinese mountain cat, E bieti
Black-footed cat, F. nigripes
Kodkod, 0. guigna
Andean mountain cat, 0. jacobitus
Flat-headed cat, P. planiceps
Fishing cat, P. viverrinus
Oncilla, L. tigrinus
African golden cat, P. aurata
Marbled cat, P. marmorata
Sand cat, F. margarita
Margay, L. wiedi
Manul, 0. manul

Chapter

1. Cats and Habitat

Vulnerability
Ranking Cat.
1
2
2
2
2
2
2
2
3
3
3
4
4
4

Table 4
Key Habitat Types for Vulnerable Cat Species
Habitat type

le
If
4a
2b
2e
2a
2c
6
Id
5a
5e

Tropical moist forest
Tropical dry forest
Grassland or shrubland
Tropical montane complexes
Second-growth
woods/fields
Tropical Savannah and woodland
Other dry woods/scrub/grass
Major wetlands
Mixed woods
High alpine and tundra
Other desert and semi-desert

Global areain 1980*(106km*)Trendover the1980s’

10.38
4.72
21.40
0.60
7.20
7.32
7.60
2.90
3.54
2.73
11.00

D
D
I
Da
I
I
S
D
Db
lb
I

Number of Species
Str-Sig AssociatedCat. I-3
Total
IO
7
7
6
6
5
5
5
4
3
3

[53%]
[37%]
[37%]
[32%]
[32%]
[26%]
[26%]
[26%]
[21%]
[16%]
[16%]

17
17
18
7
17
13
16
7
IO
3
9

* Olson et al. 1983
1 D = Decreasing; S = Stayed the same; I = Increasing.
U.S. Dept. of Energy, Oak Ridge National Laboratory,
a The tropical montane forest component
and fallow area component increased.

Lossranking system usedhere. Using a variety of literaturesources, and making extensive use of newly available
remote sensing imagery, J. Watts of the U.S. Dept. ofEnergy’s Oak Ridge National Laboratory hasupdatedthe
computerdatabase(in 1990and 1992)that forms the basisof the habitat mapsof Olson et al. (1983) (Figs. 2a-f). Dueto lack of funds, the update was not comprehensiveandhas currently ground to a halt. Quantitative data is only
available for major groupings,rather than individual habitat types (Table 8). For selectedindividual habitat types, J.
Watts (pers.comm. 1994)was able only to indicate qualitatively whether the global areahasincreased,decreased,or remainedthe samesince 1980(Table 4).
Table 8 showsthat both closed and open forests andwoodland have decreasedin global area over the past
decade. Most cat speciesare associatedwith forest, andthusmostcatshave lost a portion of their range. Watts’ figures indicate that tropical and subtropical broad-leaved
humid and dry forest has declined by 6% (+ 3%); the
FAO’s (1993) figuresindicatethat 8% is more appropriate.
The habitat type which has shown the greatest increaseover the 1980s(5% t 4%) is (3) settledareas.While nearly

Table 3
Habitat Specialists: Cats AssociatedWith Six or Fewer Habitat Types
SpeciesIssues.

Data from J. Watts,
Tennessee (pers. comm. 1994).

of this habitat type decreased,

while the highlandcrop, burnb Change very slight; could reflect an improved estimate of total area rather than actual growth or decrease.

163

[47%]
[47%]
[50%]
[19%]
[47%]
[36%]
[44%]
[19%]
[28%]
[08%]
[25%]

Part II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

However, this generally represents large-scale monoculture forest plantations. Most plantations are in the temperate regions (especially Russia, China, the U.S., Japan,
and Europe). However, tropical plantation area has been
rapidly expanding in recent years (Lanly 1982, Evans
1982, 1986, Mather 1990). Evans (1986) estimated thetotal area of forest plantations worldwide in the early to
mid- 1980s to be between 1.2- 1.4 million km2, or about 4%of closed forest area (Olson et al. 1983).
In tropical Asia, plantations of oil palm and rubber trees
are particularly significant. In Indonesia, industrial tree
plantations are likely to be the major cause of forest conversion in coming decades (Collins et al. 1991). Malaysia
is currently the top producer and exporter of palm oil andrubber in the world. Most of the production takes placein Peninsular Malaysia, where over 3 1,000 km2 of lowland
forest has been converted to industrial plantations-about
one-quarter of the original forest cover (Collins et al.
199 1). Most tropical industrial forest plantations, however, are not converted from forests but from Savannah
grasslands (which may have earlier been wooded, andhalf of the species in the family Felidae are associated withsuch areas, the degree of association is generally marginal(Table 2). The only change in habitat area which could beinterpreted as good news for cats is the increase (4% t
2%) in habitat type 4, grass and shrub complexes, withwhich a majority of both vulnerable species and the catfamily as a whole are associated. The cheetah is the best
example of a species which could strongly benefit fromthis change. However, this increase in grass- and shrubland has largely been brought about by deforestation, andintensive human modification is implicit. Under such circumstances, cats may not necessarily survive the transition (see discussion below on the factors which affect thesurvival of cats in modified habitats).
Of the 11 habitat types which harbor the greatest numbers of vulnerable cat species, five have declined in area
over the 1980s five have increased, and one is essentiallyunchanged (Table 4).
Special mention should be made of the fact that statistics from some countries seem to indicate that forest
growth and regeneration are outpacing deforestation.

Table 5
Estimates of Tropical Forest Area (in millions of km*)and Deforestation by Ecological Zone (FAO 1993)
Ecological Zone

Land
Area
(106 km*)Popufation~___I___..
~--Density
Growth(inhabitants(?4 per
/km*)
year)

Forest Cover (1990)-Area % Land Area

Deforestation (1981-l 990)
_PP.--% Forest Cover
Area

1.53

8

Forest Zone

41.86

57

2.4

17.48

1owland formations

34.86
9.47
12.89
12.49

57
41

2.3

15.44

44

1.26

Rain forest
Moist deciduous
Dry and very dry

2.2

7.18

76

0.46

Upland formations

7.01(Hi/Iand mountainforest)

42

55

2.4

5.87

70

2.3

2.38

46
19

0.61
0.22

8
6
IO
9

56

2.6

2.04

29

0.25

II

j

Non-Forest Zone
(Alpine Areas, Deserts)

5.92

15

3.1

0.08

01

0.01

IO

Total Tropics

47.78

52

2.4

17.56

37

i .54

8

Totals may not tally due to rounding.

164

Part II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

Table 6
Estimates of Tropical Forest Area (in millions of hectares’)and Deforestation by Geographic Sub-region (FAO mu)
Geographic
Sub-region/Region
AfricaWest Sahelian Africa
East Sahelian Africa
West Africa
Central Africa
Tropical Southern Africa
Insular Africa

Asia and Pacific

Number ofCountries

land
Area
(106km*)Forest Cover (10 ha*)
1980
1990

Deforestation(1981-1990)
Area Percent

40
6
9
8
6
IO
1

2,236.l
$28.0
489.7
203.8
398.3
558.I
58.2

568.6

527.6

41.0

7.2

43.7
71.4
61.5
215.5
159.3
17.1

40.8
65.5
55.6
204.1
145.9
15.8

2.9
5.9
5.9
11.4
13.4
1.3

6.6
8.3
9.6
5.3
8.4
7.6

17
6

892.1
412.2

310.6
63.9
75.2

39.0
5.5
13.2

11.2
7.9

19.3
1.1

5

190.2

5

244.4

349.6
69.4
88.4
154.7

1

45.3

37.1

135.4
36.0

Latin America and Caribbean

33

1s650.1

992.2

918.1

74,l

7.5

Central America and Mexico
Tropical South America
Caribbean

7
7
19

239.6
1,341.6
69.0

79.2
864.6

68.1
802.9

48.3

47.1

Il.-l
61.7
1.2

14.0
7.1
2.5

Totat

90

4,778.3

1,910.4

1,756.3

154.1

8.1

South Asia
Continental Southeast Asia
Insular Southeast Asia
Pacific

14.9
12.5
2.9

1 To convert hectares to km? multiply by .Ol

clearedfor agriculture and grazing) (Mather 1990).
Forestplantationsare characterizedby high timber productivity comparedwith natural forest in similar environments. In Latin America, industrial plantations make up
lessthan 1% of the forest area,but accountfor 30% of timber production. The proportion is predictedto rise to 50%
by 2000(Evans 1986). Industrial plantationscan employ alarge number of people on small areasof land and thus
relieve pressure on natural forests (Sayer et al. 1992).
However, forest plantationsare alsocharacterizedby low
biodiversity (Sawyer 1993) and, in sometemperatezone
countries(especiallyin Europe),industrialmanagedforests
have largely replacednatural forests(WCMC 1992).
The evidence on cats’ ability to makeuseof monoculture plantationsis scanty and somewhatcontradictory (seediscussionbelow). It is a subject deserving of further

research.Sawyer (1993:45)makesseveralgeneralrecommendations for plantation management to encourage
wildlife “opportunities” in industrial plantations. One
measure-allowing an understoryto developfor cover andfood for terrestrial herbivores-is specifically mentioned
ashaving benefitted the tiger in teak plantations in India(P. Jackson,pers.comm.).
Finally, statistics on habitat loss do not necessarily
include habitat degradationresulting from over-exploitation, such asover-grazing of grasslands.Degraded soils
result in a lossof vegetative productivity. This adversely
affects cats in two ways: (1) poor quality vegetation supports a lower density herbivore prey base;and (2) lossof
vegetative cover reduceshabitat suitability. According toa UNEP-sponsoredstudy of human-causedsoil degradation over the past 45 years (Oldeman et al. 1990), about

165

Part II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

Table 7
Vulnerable Cat Species Strongly or Significantly
Associated with Tropical Moist and Dry Forest
Region(s)Species

AfricaSahelian and West Africa
West and Central Africa
Tropical Southern Africa

Cheetah? leopard?
African golden cat
Lion

Asia and PacificSouth Asiac
Continental and Insular Southeast

lion

Tiger, fishing cat, rusty-spotted cat
Tiger, Bornean bay cat (Ins. only), flat-headed cat,
Clouded leopard, marbled cat, Asiatic golden cat,Fishing cat

Asia

Latin America and Caribbean
Mexico, Central America, andTropical South America

Jaguar, oncilla, margayd

a The cheetah is only marginally associated with tropical dry forest in general,but this association is stronger in the Sahel than elsewhere in Africa.
b The leopard is included here because of its virtual extirpation from the Sahel(see Species Account); tropical dry forest is a significant refuge for it in this region.
C Only those species with large or significant (e.g. fishing cat in southwestern
portions of their range in South Asia are included.
d The margay is Category 4, but is a habitat specialist
tropical moist forest.

strongly associated

10% (12.15 million km2) of the globe’s vegetated landarea(115 million km2)hassufferedmoderateto severesoil
degradationat humanhands. Most of this degradationis in
the moderatecategory; someof the soil’sability to process
nutrients into a form usableby plants hasbeendestroyed,andonly with majorimprovements-beyond the resourcesof the averagefarmer- can full productivity be restored.
Severely degradedsoils are very expensive to restore andareusually abandoned.
Soil degradationis most advani=edin Africa and Asia.
In Africa, habitat degradationhasresultedprimarily fromovergrazing by livestock, with the most affected areasinarid and semi-aridlandsin the southern,eastern,and Sahelregions. Theseregionsharbor about 55% of Africa’s 550
million head of livestock, and stock numbers haveincreasedby 75% from 1950to 1985,in spiteof the severe
droughts that occurred in most African arid zones in theearly 1970sand 1980s(Le Houerou and Gillet 1986). In
tropical Asia, soil degradation arisesmainly from deforestation, which causesmost soil degradation in South

166

India)

withAmerica as well, although not as much land has been
affected asin Asia. The Argentinian pampasgrasslands
show medium to high soil erosionfrom over-grazing andwind, and the westernslopesof the Andes show medium
to high soil erosionfrom deforestationand water run-off.
In Central andNorth America, unsuitableagriculturalpracticesaccountfor mostsoil degradation.In CentralAmerica,
only the Yucatan peninsula and the Darien Gap haveescapedthe widespreadlossof vegetative productivity.

Habitat Fragmentation
In general, loss of habitat producesa decline in species
total population size, and fragmentationof habitat can isolate smallsub-populationsfrom each other. As discussed
later in this chapter and also in Chapter 3, small discrete
populations are vulnerable to extinction due to unpredictable environmental, demographicand genetic factors,
suchas the deleteriouseffects of inbreeding depression.

Part II: Major

habitatof the South

China tiger in Sichuan,

now intensely

Chapter

1. Cats and Habitat

Lossstantially more fragmentedthan Figs. 2a-f would indicate.
While one-third of the global land surface is still wilderness,41% of it is in the Arctic or Antarctic.
Data from McCloskey and Spalding (1989) have been
modified the better to reflect non-fragmentedhabitat available to cats. Antarctica, Oceania,Greenland,and Iceland
were eliminated from the globe, as were several othercountrieswhere wild cats are not found, or occur in just asmall portion. The countries where cats are found wereassignedto the geopolitical regions usedin this Action
Plan for the speciesaccounts. The resultsare presentedin
Table 9 simply in the form of percentagesto give an ideaof regional distribution of large wildernessareas.
Analysis of the data shows that 43% of the world’s
wilderness(which totals just over 30 million km2) availableto catsoccursin the northernregionsof just two countries-Russia andCanada. Only two species,both lynxes,
occur in these areas, and then only partially, with thenorthern limit of their rangesaround the 65” N latitude.
The other major wild habitat type (20% of total wilderness)consistsof the sand and semi-desertsof Asia andnorth Africa. Desertssupport more cat speciesthan thenorthernborealforests(Table 2). However, the wilderness
criteria are weak when appliedto the Sahara,which makes
up the bulk of this wildernesstype. The region hasvery
few permanentinstallationsbut supportssome30 million
people (Le Houerou and Gillet 1986). Antelope are practically extinct in the Saharaoutsidereserves(East 1992a);the lion hasbeeneradicated,the leopardpractically so,and
only a few smallpopulationsof cheetahpersist.

Even if good quality habitat patches are within migration
distance of relatively wide-ranging species such as thecats, the intervening altered habitat can pose substantial
barriers to dispersal. Asphalt roads on which vehicles cantravel at speed, for example, have been shown to be amajor cause of mortality for small populations of cats living in fragmented habitat (e.g., Iberian lynx [Delibes
19891 and the Florida panther [Maehr et al. 199 la]). S.
Yasuma (in Zitt. 1992) has even documented several
instances of clouded leopards killed on dirt logging roads
in Kalimantan.
Global trends in habitat loss have been presented above,
and an indication of the resulting degree of fragmentation
is offered by McCloskey and Spalding’s (1989) inventoryof the world’s remaining wilderness area. McCloskey and
Spalding (1989) defined wilderness as relatively large
tracts of land (minimum 4,000 km2) free of human development, defined as man-made installations, including settlements, roads, buildings, airports, railroads, pipelines,
power lines, and reservoirs. The presence or absence ofhuman development was gauged from aeronautical maps
for pilots, which tend to show installations in remote areas
to aid navigation. Although the presence of such installations does not necessarily fragment habitat or cat populations, the methodology is more appropriate to depicting the
global state of habitat fragmentation than that used by
Olson et al. (1983), which tendsto minimize fragmentation by assigning3,000 km2 blocks of land to a predominating habitat type. Not surprisingly, McCloskey and
Spalding (1989) found that the world’s habitats are sub-FormerIssues.

cultivated.

167

Part II: Major

Issues.

ChapterI. Cats and Habitat

Loss

Table 8Percentage Change in Global Area of Major World Habitat Types Since 1980*
Global Area
in 1980(106 km2)

Percentage
ChangeNo. of Species
Cat. 1-3

Associated
Total

30.79

8% ziz3%
Decrease

16

32

10.66

SlightIncrease

5 [26%]

12 [33%]

and

5.03

5% f 3%
Decrease

6 [32%]

20[56%]

Tropical/subtropicalbroad-leaved forest (1 e & 1f)

15.10

6% iz 3%
Decrease

13 [68%]

23

[64%]

27.38

4% f 3%
Decrease’15 [79%]

30

[83%]

15.90

5%&4%
Increase

3 [16%]

17 [47%]23.90

4% f 2%
Increase

13

29

Very coJd areas withsparse vegetation (5a & 5b)

26.20

< 1%
Increase

3 [16%]

9 [25%]

Desert and semi-desert

18.20

2% t 1%
Increase

5 [26%]

16

[44%]

2.90

< 1%
Decrease

7 [37%]

14[39%]

Habitat Type

Closed forest and woodland(1

Mostly taiga and other conifer
Mid-latitude broad-leaved
mixed forest (Ic & 1 d)Open or interruptedla&woodland(2)

Settled areas (3)Grass and shrub complexes

Major wetlands(6)

1 Mainly decrease,(4(Sc-5e)lb)

but 2e (second growth woods and field mosaics) has increased* J. Watts, U.S. Dept. of Energy, Oak Ridge National

Laboratory

[84%]

[68%]

[89%]

[81%]

slightly.

(pers. comm. 1994).

several countries-Bangladesh, Cambodia, VietnamMcCloskey and Spalding (1989) indicate that no “wilderness” exists at all. While a major portion of the region
appearsasforest in Figure 2a, McCloskey and Spalding(1989)showthat it is quite fragmented. Although there arestill blocks of wild habitat remaining, none of them arevery large.
With regard to geopolitical sub-regions,habitat fragmentationis more advancedin Europethan anywhereelsein the world: 19 out of 22 countries have no wilderness,
including Spain and Portugal, home to the most endangered speciesof cat, the Iberian lynx. Central and SouthThe most species-rich geopolitical region for catstropical Asia, which covers major portionsof the rangesof
11 species-occupies only about 6% of the world’s land
area; is home to more than 60% of its people; and contains lessthan 1% of its wilderness. It is alsothe region
undergoingthe highestrate of tropical deforestation(Table5). In sharp contrast to the other regions of the world,
only 3% of tropical Asia consistsof large tracts of undeveloped land. However, it is apparentthat humanpopulation densities in this highly diverse region are such that
thereare essentiallyno large securerefugesinsulatedfrom
human activity as are found elsewherein the world. In

168

Part II: Major

America, the second most species-rich sub-region, with 10
cats, is much better off: the average national percentage
of wilderness area is relatively high at 15%, although overhalf of the region’s wilderness in found in Brazil. Overall,
60% of the 120 countries included in the data set have
< 5% of their land area as wilderness.

Implications

Chapter

1. Cats and Habitat

It is difficult to discuss the implications of habitat loss andfragmentation for cats with a great deal of confidence
because very few studies have specifically addressed thesubject. For one thing, the mechanisms by which animals
select habitats (e.g., hereditary vs. learned) is a neglectedarea of ecology (Krebs 1985) and, moreover, there has
been little research on cats in altered environments. What
may be appropriate for the few species that have been studied may not be valid for others, and any generalizations
made must be qualified by drawing attention to this lack offocused research. Most research has been, and is still, carried out in protected areas, and information on species
occurrence in altered habitats consists mainly of fortu-

RegionPercentage
of Total World
Wilderness
Area

Percentage
Wilderness
of Regional
Land Area

Percentage
Wilderness
of National
Land Area2

19%

18%

25%

16%

10%

11%

31%

14%

6%

0.6%

3%

2%

Eurasia

32%

31%

26%

5%

Americas

32%

40%

33%

I 7%

Percentage
of Total WorldLand Area1

RegionSub-SaharanAfricaNorth Africa and Southwest
Tropical

Asia

Asia

1 Total world land area considered

loss

itous sightings or anecdotal reports.
Few studies have focused rigorously on a comparisonof the ecology, behavior or population dynamics of catsin altered habitat with undisturbed habitat, with the notable
exception of Van Dyke et al. (1986a) on the behavior andmovements of a population of pumas living around an
Arizona timber concession. Other less thorough, but still
important, studies include Rabinowitz (1986) on jaguars in
a forested reserve in Belize surrounded by cattle ranches;
Seidensticker et al. (1990) on leopards at the edge of
Nepal’s Royal Chitwan National Park; Palomares et al.
(199 1) on the micro-distribution of Iberian lynx in Spain’s
Dofiana National Park and surroundings; Maehr et aZ.
(1991 b) on the movements of pumas in Florida; andZheltuchin’s ( 1992) observations on Eurasian lynx density
in logged vs. unlogged areas in Russia.
A few studies have been conducted entirely within disturbed or modified habitat: Bowland’s (1990) study of servals on farmland in South Africa; studies of leopards on amixed livestock/game ranch in Kenya (Mizutani 1993) andin a mountainous farming region in South Africa (Nortonand Lawson 1985); and studies of jaguars on cattle ranches
in Brazil (Schaller and Crawshaw 1980, Crawshaw andfor Cat Species

Table 9Wilderness Area by Geopolitical

Issues.

as just over 120 million km*.

2 Average of the wilderness area percentages for each country in the region. Sub-SaharanAfrica: n=40 countries;
North Africa and Southwest Asia: n=19 countries; Tropical Asia: n=lO countries; Eurasia: n=28 countries;
Americas: n=23 countries.
/Vote: Data from McCloskey and Spalding (1989) have been modified the better to reflect non-fragmented
habitatavailable to cats. Antarctica, Oceania, Greenland, and Iceland were eliminated from the globe, as were several othercountries where wild cats are not found, or occur in just a small portion.

169

Part II: Major

Issues.

Chapter

1. Cats and Habitat

Lossconsidered to be primarily nocturnal or crepuscular in
India, but after they were strictly protected in somereserves, such as Ranthambhore National Park, daylightactivity was frequently observed (Thapar 1992).
Some degree of habitat fragmentation is demonstrably
beneficial to cats. The two habitat types with the most cat
species associated (28 species) are both fragmented habitat
types: 2e, second growth woods and field mosaics, and4a, grass and shrubland (Table 2). These two types ofhabitat often arise from human activities. A mosaic oftrees or shrubs and grassy vegetation may be ideal for mostcats. This is related to the fact that edge environments aregood for hunting (e.g., Eaton 1970, Sunquist 1981, Prins
and Iason 1989), for two reasons: (1) clearings and forest
gaps are good habitat for both small and large herbivorous prey species, primary productivity being greater thanin closed forest (e.g., Leopold 1933) and; (2) cats mainly
hunt by sight, and an edge environment gives a cat cover tohide in and a view over which to spot prey. Two North
American cats for which habitat use has been well studied-the Canada lynx (intermediate habitat selectivity)and bobcat (broad habitat selectivity)-havebeen found tohunt mainly in forest clearings made by fire, logging,
farms, and roads (Hall and Newsom 1978, Miller 1980,
Parker 1981).
Panwar (1987) and McNeely (1994) have pointed outthat traditional systems of shifting cultivation, which haveexisted for over 10,000 years in tropical Asia, make forgood habitat which supports a high diversity of wildlife.
Large ungulates feed in the abandoned fields, providing
prey for the bigger cats. Older fields contain a high proportion of fruit trees on which a variety of smaller arboreal animals and birds come to feed, providing prey forthe smaller cats. Closed forest patches provide the catswith shelter. Wharton (1968) has provided strong evidence that the distribution of the major large mammals ofsoutheast Asia is highly dependent on shifting cultivation,
because mature tropical forests conceal most of their edible products high in the canopy beyond the reach of terrestrial herbivores, while clearings bring the forest’s
productivity down to where it can be reached. Karanth andSunquist (1992) reached a similar conclusion regarding thedensity of herbivores in mixed habitats, noting that maintenance of clearings with successional vegetation within
the tropical forest of southwestern India’s Nagarhole
National Park was at least partly responsible for high herbivore biomass, of an order comparable to or higher thanthat found in African Savannah parks. Nagarhole also supports relatively high tiger densities (Karanth 199 1).
Modified habitats that are considerably less optimal
than tree and grass mosaics can still be used. Pumas were
only rarely located in or moved through clear-cut logged
areas in Arizona, but one female established a home range
which included such an area. Although she used both the

Quigley 1991). Others have examined the diets of big catsnear livestock ranches; the findings of these studies are
addressed in the next chapter.
Although habitat loss is often described as the primary
threat to cat populations, there are several indications thatcats adjust relatively well to many forms of habitat loss
and fragmentation, including deforestation, with only
extremes such as urban settlement being generally devoidof cats.
Most cats make use of a variety of habitats, and thisnot only buffers them against the loss of a preferred habitat type, but also suggests substantial flexibility in habitat
selection and use. Based on studies of the bigger cats,
Sunquist and Sunquist (1989) concluded that suitabilityof habitat for the bigger cats is determined primarily byavailability of adequate amounts of cover and prey; availability of adequate water could also be added. The index
of species-habitat association used here agrees: cats occur
in an average of 7.5 broadly defined habitat types. The
generalist label does not include the entire family: 39% of
the cat species were identified as having narrow habitat
selectivity (Table 3). However, species identified as having intermediate to broad selectivity are among the betterstudied, while of those identified as specialists, only for
one species, the Iberian lynx (with the smallest contemporary range of any cat species), has habitat association
been investigated specifically (Palomares et al. 199 1).
There have been reports of some of these specialist species(kodkod, flat-headed cat, marbled cat) occurring inhuman-altered environments, including farms, forest plantations, and logged forest (see Species Accounts).
At the same time, cats exhibit flexibility by behaving
differently in disturbed and undisturbed habitats. Pumas
tend to be inactive during the middle of the day, more
active during some mid-evening hours and sunset, andmost active at or just prior to sunrise (Ackerman 1982,
Van Dyke 1983). Van Dyke et al. (1986a) found thatpumas in the vicinity of logging concessions and humandisturbance shifted activity peaks to after sunset, concentrated other activity during night hours, and were inactive
rather than active at sunrise. This pattern held true both
between different pumas living in disturbed and undisturbed habitats, and for the same individual whose homerange included both types of area. ‘Others have reachedsimilar conclusions on the basis of more anecdotal evidence. Geertsema (1985) reported that servals are almost
entirely nocturnal (rather than crepuscular) in areas of highhuman activity, even though some of their main rodent
prey are predominantly diurnal, and prey densities can bequite high around agricultural areas (Smithers 1978). In
Sumatra, Griffiths and van Schaik (1993a) similarly
reported that tigers in Gunung Leuser National Park wereprimarily diurnal or crepuscular, but almost exclusively
nocturnal close to human settlement. Tigers were long170

Part II: Major

logged area and other less disturbed areas, the logged area
was seldom visited during periods of human activity (Van
Dyke et al. 1986a). Van Dyke et al. (1986a) attributed
the reluctance of pumas to use logged areas primarily toloss of cover, while pointing out that increased humanactivity and road density were also deterrents.
Johns ( 1986, 1989, 1992) has examined the effects oftropical forest logging on wildlife, and has concluded that“few predators are eliminated by logging operations inrain forest” (Johns 1992:45). In the tropics, logging is
usually not as destructive as in the temperate zone. Whiledamage associated with the “selective” harvesting of commercially valuable species can leave up to 30-40% of aconcession bare of vegetation (Kartawinata
1978), thiscompares favorably with the results of temperate zone
clear-cutting as practiced in the Arizona study area
referred to earlier. In general, Johns (1989) suggests thatfew mammals are specialized exclusively for mature forest vegetation. Tree falls and fires create both small andlarge gaps in forests, and subsequent secondary regeneration is an essential part of forest cycles (Pickett et al.
1992). Johns (1989) recorded both clouded leopard andmarbled cat in six-year-old logged forest in Sabah, andsmall cats (unidentified) were seen in logged forest in
Peninsular Malaysia every year from one to 12 years after
logging. While admitting the difficulty of sorting out
whether species occur in logged areas only due to availability of unlogged blocks, isolates, or riverine refuges,
Johns (1989:74) speculates that “large predators, such as
clouded leopards...may be able to persist solely within
logged forest if the area is large enough.” However, in
Peninsular Malaysia, he found that tigers and leopards
were largely absent even from older logged forest, despite
the availability of prey species-including
deer, mousedeer, and seladang or gaur-whichwere attracted to thegrasses and regenerating growth. He concludes that theirabsence was probably due directly to persecution fromhunters subsequent to the lifting of access restrictions by
the logging company.
Sunquist and Sunquist’s (1989) characterization of
habitat suitability mentioned above can also be stated inthis way: the only type of habitats not suitable for cats arethose which have inadequate or unsuitable cover and prey(and water). The main exception would be a case where
habitat alteration benefits one cat species to the detrimentof another through interspecific competition. The clearing of coniferous forests for agriculture in the northern
United States is one example: this has permitted the bobcat
to expand some parts of its northern range, while the southern boundary of Canada lynx range has retreated (Rolley1987). Although the physical structure of human-alteredhabitat is changed-sometimes
radically-alternative
sources of food and cover are generally available.
Populations of rodents and birds, different from those

Issues.

Chapter

1. Cats and Habitat

Lossfound in forests, thrive in croplands and even cities. When
forest is cleared for pastureland, wild ungulates may bereplaced by domesticated livestock.
When settlement
encroaches on woodland and ungulates become scarce,
pumas, leopards, tigers, and snow leopards have been
known to take dogs. That rusty-spotted cats, a little-known
species with a restricted distribution in India, have beenfound on several occasions to den and breed in the ceilingspace of houses, while preying on domestic poultry (seespecies account), is testimony to felid adaptability.
However, the definition of “inadequate” and “unsuitable” will obviously vary among species, and it would be
misleading to conclude that all cats are generalists thatcan live anywhere and eat anything. For example, somecats may be constrained in their ability to live in certain
habitats through specialization for a certain size categoryof prey. If that size category of prey were to disappear(e.g., because of overhunting by humans) and were notreplaced by similar species, cats might not be able to survive even if other, different sized prey, were available.
Breitenmoser and Breitenmoser-Wiirsten(1990) havediscussed this subject with respect to the 20th century
extirpation of the Eurasian lynx from southern Europe.
The Eurasian lynx, the largest of the lynxes, is adapted toprey chiefly on smaller ungulates (unlike the three otherlynxes, which favor rabbits). In southern Europe, small
ungulates consisted of two species, the roe deer and thechamois, until they were almost completely extirpated by
the late 19th century. Although livestock (sheep and cattle) were available, husbandry in the mountain uplands ofthis region tends to be migratory, whereas the lynx, likemost cats, is territorial. While direct persecution may havebeen the primary cause of the extinction of lynx in the lowlands, an inadequate prey base in the highlands, despite
good quality habitat and availability of small prey, appears
to have been a major factor in the loss of the lynx.
Inadequate suitable prey could also explain the disappearance of other cat species from viable habitat, such as
the lion and leopard from North Africa, where the wild
ungulates have been severely reduced in number, and thelivestock husbandry which has replaced them is nomadic.
Of the big cats, only the cheetah, which follows migrating
herds of Thomson’s gazelle in the Serengeti (females andnon-resident males only: Durant et al. 1988), has persisted
in any number in the Sahara and Sahel regions. Similarly,
the rarity of the Andean mountain cat may be attributable
to the virtual extinction of the mountain chinchilla (seespecies account).
This review indicates that habitat loss and fragmentation
are not the main threat facing cats, although they are closely
and inextricably linked to it. Degradation of the vegetation base upon which herbivores depend may reduce thenumber of suitably-sized ungulate prey so that cats cannot
sustain themselves. Although shifting cultivation has been

171

Pat? II: Major

Issues.

Chapter

1. Cats and Habitat

Losscharacterized as good wildlife habitat, this is only true if
carried out in a traditional, sustainable manner. Otherwise,
severe habitat degradation can result (McNeely 1994).
Persecution, both direct and indirect, is the primarycause of declining cat populations. Indirect persecution
consists chiefly of overhunting the prey species uponwhich cats depend. If the only suitable prey available inmodified habitat is livestock, persecution then becomes
direct. This problem is addressed in the next chapter,
Management. The most direct form of persecution is commercial hunting, which is discussed in Chapter 4, Trade.
It is not that conservation of natural habitat is unimportant for cats: on the contrary, it is because of losses topersecution that the need for relatively undisturbed conservation areas becomes paramount. Rather, it is a matterof emphasis and prioritizing limited resources for cat conservation. Cats are wide-ranging and thinly distributed,
and for most it will be impossible to protect all but a small
proportion of their historic ranges. It is encouraging thatcats are among the more adaptable species which are capable of coping rather well with the changes that the growing
human population is making to the world environment.
Human settlement, activity, and development are not necessarily inimical to cats. Future efforts need to be directed
towards understanding to what degree cat species tolerate
habitat loss, fragmentation, and modification; developing
guidelines as to how land use can be modified to allow cats
to persist; and finding ways to reduce persecution of cats
(especially the big cats) in habitat used by humans. The
following review of the world’s system of protected areas
and their significance for cats will show why the issue ofconservation in human-occupied landscapes is so critical.

is encouraging that the regions with the greatest proportionof protected land area are in the tropics. About 5% of thetropical forest biome is now protected in reserves (WCMC
1992). Heywood and Stuart (1992) have noted that thispercentage is rising rapidly, and has its distribution heavily
skewed towards the most diverse sites (as indicated by bird
and mammal distributions). While legally protecting habitat has had only a minor impact on global deforestation
rates, there have been important advances in mitigating the
anticipated effects on biodiversity.
No cat species lack protected areas within their ranges,
and the presence of all but two (Chinese mountain cat,Andean mountain cat) has been confirmed in at least one
protected area. No cat species are presently confined only
to protected areas.
Ehrlich’s (1988) assertion that “the loss of geneticallydistinct populations within species is, at the moment, atleast as important a problem as the loss of entire species” ishighly relevant to the widely distributed cat family.
Overall, regional populations or subspecies are more critically threatened than species, and the most important contribution that protected areas currently make to cat
conservation is helping to prevent intraspecific genetic
erosion by protecting important sub-populations.
The tiger is a good example of how quickly local extirpation can happen. Two distinct island populations (Baliand Java) and an isolated race found along watercourses in
the west Asian deserts (Caspian tiger) have been lost
within the past 20-40 years. A fourth subspecies, theSouth China tiger (which, based on morphological evidence, appears to be the most primitive tiger from which
the other races evolved [Herrington 1986]), has been
reduced to a handful of scattered individuals. The Amurtiger is seriously threatened by a very recent wave of commercial poaching, and the Sumatran race, confined mainly
to the island’s major reserves, is estimated to number only
400-600 (Tilson 1992a).
Using the snow leopard as another example, two subspecies have been named but are not generally recognized(Hemmer 1972). A. Kitchener (in Zitt. 1993) has suggested
that the species is a prime candidate for significant
intraspecific variation, given the isolating effects of boththe permanent snowfields at high altitude and densely settled valleys within its range. With a patchy distribution,snow leopards are particularly vulnerable to local extirpation, which has been anecdotally reported across much of
their range (Jackson 1992).
While tigers are increasingly rare outside protectedareas, most snow leopards inhabit unprotected lands. The
different situations of these two species, the rarest of the
big cats, each with an estimated global population ofaround 6,000 (Jackson 1993a, Fox 1994), make usefulcase studies to illuminate both the benefits and limitationsof the role currently played by protected areas in cat con-

The Role of Protected Areas
in Cat Conservationand theNeed for Linkages
According to WCMC (1992), there are more than 8,800
protected areas in 169 countries covering some 7.7 million
km* or 5 .2% of the earth’s land area (not including multiple-use or partially protected hunting areas, nor private
reserves.) Most of these have been established relatively
recently, i.e. since the early 1960s. Table 10 outlines theprotected area network of this Action Plan’s five geopolitical regions, based on data published by the World
Conservation Monitoring Centre. Although the figuresmask a good deal of variation between countries, broad
regional trends are apparent. In Africa and southwest Asia,
protected areas tend to be few but large, while in Eurasia,
they are many but small. Tropical Asia and the Americas
fall somewhere in between.
Given the prevailing rates of tropical deforestation, it

172

Pat? II: Major

Table 10
Protected Area by Geopolitical
RegionIssues.

Chapter

1. Cats and Habitat

RegionPercentage of Land Area Protected

Number of Protected Areas

Regional

Average

Regional

Average
National

Average
Size (km2)

5.2%

5.2%
O-17.4%
4.55

599

15.0

1,989

3.4%

1.9%
O-l 0%
3.0

230

12.1

2,056

2.7%

7.9%
0.1925%
6.3

3,454

123.4

306

Tropical Asia
Range
Standard deviation

5.9%

6.3%
O-I 9.5
5.7

865

86.5

519

Americas
Range
Standard

7,0%

7.0%
o-31 %
7.2

2,189

95.2

1,229

Sub-SaharanAfricaRange
Standard deviation
North Africa and Southwest
Range
Standard deviation
Eurasia
Range
StandardAsia

deviationLossdeviation

Source: WCMC (I 992)

servation. SinceIndia’s approachto tiger conservationhas
not only beenpioneering,but hasalsorelied mainly upon
protected areas,discussioncenters largely on that country. For the snow leopard, the role of protected areasin
conserving minimum populations has been examinedrecently by Jackson and Ahlborn (1990), Fox (1994),
Green ( 1994),and Villarubia and Jackson( 1994).
Seriouspublic concern about the decline of the tiger
aroseonly in 1969at the 10thsessionof the IUCN General
Assembly in New Delhi, where a precipitous decline in
numbersof the Bengal tiger was documented. Seven of
the eight tiger subspecies
were already listed in the IUCNRed List of Threatened Species, and as a result of theAssembly discussions,the Bengal tiger joined them. The
Indian government subsequentlyplaced the tiger under
total legal protection from hunting and trade, and conducted a nationwide census. The resulting estimatewasof the order of 1,800 tigers (Govt. of India 1972), comparedwith a possible40,000 at the turn of the century (Gee1964). Project Tiger was launchedby the governmentin
1973, with the objective of employing the tiger asa flagship speciesto focus effort on habitat preservation andconservationof sympatric species.Both central and state
governmentresourceswere concentratedon improving theviability of a set of “Tiger Reserves” selectedaccording
to ecologicaldiversity (23 in 1995). Annual expenditures
have beenof the order of U.S. $4,000,000, and the states
have sustainedan annuallossof about$14,000,000by giving up logging operationswithin reserveborders(Panwar
1987). The reserveswere designedwith central core areasafforded total protection, surrounded by buffer zones
where somelocal exploitation, suchaslivestock grazing ortree felling, may be permitted. The government cametorecognize that conservation-orientedcommunity development in the areassurroundingthe Tiger Reserveswould bevital to Project Tiger’s success(Panwar 1987, Anon.
199Of). However, in denselysettledand poverty-stricken
rural India, reconciling the needsof nature reserveswith
thoseof surroundingcommunitieshasproved an elusive
goal (Thapar 1992, Govt. of India 1993, Jackson1993b),
andprogressin implementationhasbeenslow andpatchy.
Overall, Project Tiger has been successfulin its primary objective of conservingthe tiger; the decline wasnot
only arrested,but reversed. According to the resultsof the173Part II: Major

Issues.

Chapter

1. Cats and Habitat

Dolomite

Loss

mines

infringing

on Sariska

Tiger Reservein India.

but the unreliability of the census figures make an accurate
assessment impossible. Moreover, seven Tiger Reserves
have become refuges for political militants, which has
made management “exceedingly difficult” (Govt. of India
1993). Most of the reserves are subject to illegal grazing
and tree felling, and all suffer from the antagonism of people living just outside reserve borders. Thapar ( 1992) has
documented the destructive nature of a system of complicity and corruption between state-assigned officials andvillagers in the area of the Ranthambhore Tiger Reserve,
leading to severe habitat degradation and a breakdown of
law and order.
The situation of the snow leopard is similar, but not as
drastic, given the difference in human population densities between the central Asian highlands and southern
Asian lowlands. Green (1994) reviewed the status of 47
protected areas in the mountains of central Asia where
snow leopard presence has been confirmed. Of these, 65%
are inhabited by people, and 86% provide grazing for livestock. Livestock predation is a major problem (Miller andJackson 1994), and commercial poaching for pelts andbones has been reported (R. Jackson, pers. comm.). In
general, there has not been much effort to integrate local
requirements with conservation objectives, although there
has been progress in Pakistan (Ahmad 1994) and Nepal (R.
Jackson, pers. comm.).
Similar programs are being
planned in China (R. Jackson, pers. comm.) and India(Govt. of India 1988).
These problems are not unique to India, a densely populated country, nor to central Asia, where populations are

1993 nationwide census, the tiger population was officially estimated to be 3,750 (Nath 1994), more than double the 1972 figure. There have been numerous othergains, as well. The tiger effectively served as an umbrella
species, and other species have benefitted from improved
habitat protection and management in reserves with tiger
populations. Perhaps most importantly, the tiger served asa charismatic flagship species around which the conservation movement in India rallied and grew. When Project
Tiger was initiated, government forests (which make up
most of India’s forest cover: Govt. of India I99 1) weremanaged almost exclusively for timber production; but
thereafter the importance of wildlife conservation was recognized. In the Project’s early years, there was a pervasive atmosphere of enthusiasm and hope among theconservation community, especially the government officials charged with the tiger’s conservation(M.K.
Ranjitsinh, pers. comm.).
However, the main threats posed to the gains made by
Project Tiger indicate the first important obstacle to realizing the full potential contribution that protected areas canmake to the conservation of cat populations: unless theresource needs of local people are somehow satisfied andintegrated into reserve management, no officially protected area can really be protected effectively on the
ground. The Indian government acknowledges that commercial poaching for tiger bone has become a serious problem in the 1990s (Govt. of India 1993). The difference
between the 1993 census figure of 3,750 and the 1989 figure of 4,334 suggests that nearly 600 tigers have been lost,174

Part II: Major

Chapter

1. Cats and Habitat

Losshostile environment need to be large enough to protect
against the extinction pressures which threaten small populations. The concept of minimum viable population(MVP) size is discussed more fully in Chapter 3, Research.
It should just be noted here that an MVP is a relatively
small, isolated population, but one still large enough sothat the chances are good that it will survive for at least 100
years. Extinction pressures which threaten its survival
include unpredictable environmental and demographic
events (e.g., a virulent epidemic, a massive forest fire, or
disruption of breeding dynamics, such as through a chance
imbalance of sex ratio), in addition to harmful genetic
processes (accumulation of deleterious mutations, loss ofgenetic diversity) to which small populations are vulnerable. As a general rule, an isolated MVP should number inthe hundreds of breediltg individuals-whichmeans
potentially a thousand or more animals (Lande andBarrowclough 1987).
With reference to field conservation, small population
biology is still largely a theoretical rather than an appliedscience (Caughley 1994). Although there are questions
regarding the consequences to wild populations of loss ofgenetic diversity (see, for example, the cheetah Species
Account), prudence dictates that the warnings of small
population biologists be heeded.
The vast majority of the world’s protected areas are tooof much lower density but clustered. Protected areas
around the world face the same sort of threats emanatingfrom outside their borders, and managers are increasingly
focusing on developing locally appropriate solutions toresolve conflicts between parks and local people (McNeely
and Miller 1984, MacKinnon et al. 1986, IUCN 1992b).
The privately-sponsored
Ranthambore Foundation inIndia is relevant as a case study. It was established in 1987
to support development projects aimed at reducing villages’ dependence on the Ranthambore Tiger Reserve,
which is coming under increasing pressure as the land outside the reserve has suffered severe degradation. The
Foundation’s activities include distribution of indigenous
seeds for tree planting, programs to improve livestock
management, provision of primary health care and promotion of innovative and sustainable income generation
projects, The Foundation also supports educational andcultural activities aimed at restoring traditional links to andrespect for the Ranthambore environment (Fig. 3). International financial support is provided by an affiliated organization, Global Tiger Patrol, a British-based charity
which organizes and supports similar programs elsewherein tiger range.
The second major problem with protected areas forcats is that big cats, in particular, are wide-ranging andthinly distributed, and isolated reserves surrounded by aRanthambhore
of a program

Issues.

Foundation
medical service for villages around tiger reserve,
Ranthambhore,
to improve the lives of local people and relieve pressure on tiger habitat.

175

India, partPart II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

production (H. Panwar, pers. comm. 1988).
The snow leopard’s status in Nepal resembles that of
the tiger in India-no protected areas are large enough to
support viable breeding populations (Fox 1994)-but,
unlike the tiger, most of its good quality habitat lies outside
the reserve network (Jackson and Ahlborn 1990, Fox
1994). Assuming snow leopard presence in all suitable
habitat types, and using a range of densities according to
the degree of habitat suitability, Jackson and Ahlborn(1990) estimated that 65% of Nepal’s snow leopard population occurs outside protected areas. Out of all the country’s national parks and reserves, only the proposed
Annapurna Conservation Area-if
fully implementedwould come close to harboring a large population (projected at 156 adults based on habitat suitability). However,most villagers surveyed who live within one part of theAnnapurna CA wished to see snow leopards completely
eradicated because of livestock predation (Oli et al. 1994).
The tiger population in the Indian sub-continent ishighly fragmented. Smith andMcDougal(l99 1) estimatedthat most sub-populations have fewer than 10 breeding
individuals, with very few ashigh as50. Tiger populationsin Thailand are alsoknown to be widely separated.Snow
leopard populations, on the other hand, tend to be naturally fragmented due to the patchy nature of high alpine
habitat, dissectedby deepriver valleys which are heavily
settled (Jacksonand Ahlbom 1990,Fox 1994). For mosttiger populations,there is no effective movementbetween

small to support MVPs, certainly of the bigger cats, and ofmany of smaller species as well. Approximately one-thirdof the world’s protected areas are between lo-29 km2 insize, and 85% are less than 1,000 km2 (WCMC 1992).
To return to the Indian tiger: most of the 21 Tiger
Reserves which are managed by Project Tiger are virtual
islands of habitat in densely settled areas and, even including their partially protected and often highly disturbed
buffer zones, they may not be large enough to conserve
viable tiger populations.
The areas of the 21 Tiger
Reserves in India range from 521 km2 to 3,568 km2 andaverage 1,500 km”, but more than half are less than 1,000
km2 in size. According to the 1993 census, based on pugmarks, the average tiger population in each reserve was 65(range 17-25 1), including sub-adults. Yet these censuseshave been criticized as unreliable because of imprecise
methodology (Karanth 1987, 1993b), and poaching has
severely reduced the population in at least one reserve (V.
Thapar, pers. comrn.). It is thus likely that breeding populations within these reserves are considerably lower thanthe official estimates would indicate. Tigers are also foundin approximately 100 other protected areas in India, but
the average size of these reserves is less than half as large as
the Tiger Reserves (Johnsingh et al. 199 1; Wildlife Instituteof India unpubl. data). Strictly protected areas make up
only a small part of India’s forest cover. According to census data, more than half of the country’s total tiger population lives outside these areas, in forests managed for timber

Tigers view devastated
Painting by Gajanandland around Ranthambhore
Tiger Reserve
Singh, Ranthambhore
School of Art.

176

which was still forest

in the 1960s.

Par? II: Major

Issues.

Chapter

1. Cats and Habitat

Loss

The appropriate way to proceed would seem to be torecognize the potential significance of corridors for cat
conservation, but to judge each case according to its merits. Beier (1993) and Maehr (1990) both document theimportance of short, narrow natural landscape features infacilitating movement between fragmented puma sub-populations in southern California and southern Florida.
These are clearly cases where maintenance or restorationof corridors would benefit locally scarce cat populations.
Research effort should increasingly be directed towards
examining use of modified habitat by cats, and to identifying key habitats and potential corridors in settled areas
where cat populations are either clustered or locally scarce.
A GIS computer-based map overlay system is an asset tothis process (Smith et al. 1987a).
In many cases, habitat corridors used by cats will beon privately owned land. For example, Maehr (1990) has
emphasized that many pumas in Florida occur on private
land, and that larger blocks of protected land are of marginal quality and support fewer animals. Governments
should explore options beyond outright acquisition of privately owned wildlife habitat for protected areas. The best
options provide economic incentives to land-owners tomanage their land in a way compatible with species conservation. In developed areas, these would include bothcase-specific legal agreements as well as a broader system of tax write-offs (e.g., Logan et al. 1993). In developing areas, this could either take the form ofpreferentially focusing development activities on lands
supporting key wildlife habitats or, as has been pioneeredin southern Africa (Martin 1986), helping land owners todevelop the knowledge and infrastructure to make maintenance of wild lands a type of land-use which is economically competitive
with far more prevalent (anddestructive) agriculture or livestock grazing.
There is a third reason why it is important for cat conservation to look beyond strictly protected areas: there are
a number of constraints which limit the amount of habitatthat governments can set aside as strictly protected. At
the present time, protected areas comprise only a minor
portion of cat species’ ranges. If cat populations are lost
over the next decade, they will most likely be lost on
unprotected lands.
An analysis of the representation of cats in the world’s
protected areas has been carried out using data collected by
the Protected Areas Data Unit (PADU) of the World
Conservation Monitoring Centre. The figures presented inTable 11 are preliminary and necessarily inaccurate: dataon confirmed occurrence of cats in protected areas is neither sufficiently comprehensive nor centralized, so that itwas necessary to assume that protected areas falling withina species range actually contain the species (see Appendix
4 for details on methodology).
Rabinowitz’s (1993) survey of tiger occurrence in pro-

populations. The degree of migration between snow leopard populations has scarcely been examined, but is probably impeded in many places by settlement. Even a very
low rate of immigration can, theoretically, substantially
improve the viability of even very small populations(Allendorf 1983, Lande and Barrowclough
1987, Beier
1993). Beier’s (1993) population viability analysis forpumas in southern California and Florida lends weight toa growing advocacy within the conservation community of
the use of “corridors” to connect protected areas and permit
movement between sub-populations.
Beier ( 1993) developed a density-dependent model thatsimulated the dynamics of small populations of pumas
under varying environmental and demographic conditions.
He did not consider the effects of inbreeding or genetic
drift-taking
genetic considerations into account greatly
increases MVP size (Seal et al. 1989). His model indicatesthat if a wildlife movement corridor is available to allow
immigration of just a few individuals per decade, an area
as small as 600- 1,600 km2 (depending on the demographics of the particular population) can support a puma population without significant extinction risk for 100 years.
Isolated populations without immigration had a muchhigher risk of extinction, and did not persist “safely” inareas under 2,200 km2. Beier’s study (Beier and Barrett
1990- 1992) of a small puma population (about 20 adults)
in the Santa Ana Mountains of southern California documented the use of narrow corridors (one along a creek andanother through a canyon) for both dispersal and movement between larger habitat blocks.
Corridors have attracted much attention in the wake ofgrowing awareness of the vulnerability of small isolated
populations, and because the concept of linkages between
larger blocks of habitat makes sense intuitively(Simberloff et al. 1992). A corridor strategy, according
to one of its chief proponents (Noss 1987, 1991), consistsof maintaining or restoring natural landscape connectivity,
not building connections between naturally isolated habitats. Because of their range requirements, large carnivores,
particularly cats, are often used as examples of species
which would benefit from corridors. Soule (1991), pointing out that large animals, particularly predators, arethought to play a keystone role in maintaining habitat andspecies diversity, deems it “ecologically prudent” toattempt to ensure their presence, through corridors, insmall habitat patches which could not otherwise support apermanent population. However, corridor strategies havebeen criticized because of the potential expense of purchasing narrow pieces of land in the vicinity of settlement
or development (the same amount of money could buy
more land in more remote areas). Critics further point toa lack of species-specific research documenting use vs.
non-use of habitat-corridors (Simberloff and Cox 1987,
Simberloff et al. 1992).
.
177

Part II: Major

Issues.

ChapterTable 11
Protected

1. Cats and Habitat

Loss

Area for Cat Species (in order of vulnerability)

Species% of Range
Protected

Amount of Area
Protected (km2)

Number ofProtected
Areas25-27%

1 o,ooo-50,000

50-I 50203%12.14%
9-l 2%
25-27%
6-9%
6-9%
36%
6.9%
12014%
6-9%

1 o,ooo-50,000
1 o,ooo-50,000
1 OO,OOO-250,000
1 o,ooo-50,000
1 o,ooo-50,000
1OO,OOO-250,000
1 OO,OOO-250,000
1OO,OOO-250,000
1 OO,OOO-250,000
1OO,OOO-250,000

1 o-50
1O-50
1 O-50
1 O-50I O-5050-I 50
290-350
50-I 50
290-350
50-I 50

6-9%
9-I 2%
12-14%
396%
9-12%
9-I 2%
306%
6-9%

250,000-500,000
250,000-500,000
250,000-500,000
I o,ooo-50,000
250,000-500,000
550,000-700,000
250,000-500,000
150,000-250,000

150-260
600-750150-260
50-I 50
600-750
290-350150-260
290-350

3-6%
6.9%
6-9%
12-l 4%
2-3%
3-6%

1OO,OOO-250,000
550,000-700,000
550,000-700,000
550,000-700,000
1 o,ooo-50,000
1 OO,OOO-250,000

1 O-50290-350
290-350
400-500
50-I 50150-260

306%
203%
306%
306%
6.9%
6-9%

550,000-700,000
250,000-365,000
250,000-365,000
1OO,OOO-250,000
I-l .7 million
l-l .7 million

400-500
400-500
600-750
290-350I OOO-1,500
1000-1,500

3-6%
3-6%
3-6%

I-l .7 million
250,000-500,000
250,000-500,000

600-750
400-500
IOOO-1,500

3-6%
306%

I-l .7 million
550,000-700,000

IOOO-1,500
400-500

Category 1
lberian lynx, L. pardinus

Category 2
Bornean bay cat, C. badia
Chinese mountain cat, F. bieti
Black-footed cat, F. nigripes
Kodkod, 0. guigna
Andean mountain cat, 0. jacobitus
Flat-headed cat, P. planiceps
Fishing cat, P. viverrinus
African golden cat, P. aurata
Tiger (A), P. tigris
Snow leopard (A), U. uncia

Category 3
Cheetah (A), A. jubatus
Asiatic golden cat, C. temmincki
Oncilla, L. tigrinus
Rusty-spotted cat, P. rubiginosus
Clouded leopard (A), A/. nebulosa
Lion (A), P. lea
Jaguar (A), P. onca
Marbled cat, P. marmorata

Category 4
Sand cat, F. margarita
Margay, L. wiedi
Serval, L. serval
Canada lynx, L. canadensis
Geoff roy’s cat, 0. geoffroyi
Manul, 0. manul

Category 5a
Ocelot, L. pardalis
Eurasian lynx, L, lynx
Bobcat, L. rufus
Pampas cat, 0. colocolo
Puma (A), P. concolor
Leopard (A), P. pardus

Category 5b
Caracal, C. caracal
Jungle cat, F. chaus
Leopard cat, P. bengalensis

Category 5c
Wildcat, F. silvestris
Jaguarundi, H. yaguarondi

See the Introduction

to the Species

Accounts

for explanationof the vulnerability

178

rankingsystem(pp. Z-6).

Fart II: Major

tected areas in Thailand shows the risks of this approach:
only 58% (n=22) of surveyed protected areas contained
tigers, and the six largest sites contained 56% of the total
estimated number of tigers in Thailand. Nonetheless, thefinancial incentive to poach tigers is high. It is this species
that is most likely of the cats to be eliminated from protected areas, and it is for this species that the assumption of
occurrence is the weakest.
Table 11 shows that, even for a species which has disappeared from much of its former range and now exists inhighly fragmented sub-populations, the Iberian lynx, protected areas comprise only about one-quarter of its range.
Most species have less than 9% of their range protected.
Most of the cats with a greater proportion protected arevulnerable species ranked in Categories l-3. Species withsmaller ranges are more likely to have a greater proportion of their ranges protected than species with larger
ranges, although the actual geographic area protected issmaller. The more unprotected range a cat loses, the more
important protected areas become.
Protected areas are necessary for cat conservation and,given prevailing rates of habitat loss and fragmentation,
their importance is likely to increase with time. However,cat conservation must have a two-pronged, interactive
strategy. Protected area coverage for cats should be
improved by: (1) establishing new protected areas to conserve important habitat or populations; (2) strengthening
the protective infrastructure of threatened areas; (3) generating local community support for maintaining the protected area; and (4) taking measures to ensure that
protected populations are of viable size. Furthermore, inorder to better conserve both inter- and intraspecific diversity, as well as to promote connectivity between populations in protected areas, efforts must also be directed
toward conserving cats in places used more intensively
by people. This is especially problematic for big cats, as
discussed in the next chapter.

Summary
4

Issues.

Chapter

7. Cats and Habitat

Lossregional habitat type. Habitat conservation for cats isa priority here.
2. Four habitat types are of greatest importance for vulnerable cats: tropical rain forest, major wetlands, tropical montane complexes, and high alpine tundra. The
first three types are declining in global area; the latter
three habitats share the features of being small in overall area, and locally patchy or insular.
3. Global trends in habitat loss should not provoke undue
pessimism as far as cat conservation is concerned. Cats
appear to be relatively flexible in their habitat requirements, and can persist in many types of modified habitat. Research effort needs to be directed at defining thecommon denominators which allow such persistence(e.g., Projects 2 and 23 in the Action Plan, Part III).
4. No cat species appears to be imminently threatenedwith extinction due to habitat loss, but significant
threatened sub-populations may need immediate protection through reserve creation. Status surveys should
be conducted so that such populations can be identified;a number of priority projects in Part III are aimed at
this. It is imperative that the establishment of newreserves closely involve local residents so that theeffectiveness of the protected areas is not jeopardized.
5. The protected area network plays an important role inconserving individual cat populations. However, if
isolated, most reserves are too small to support minimum viable populations.
Corridors which permit
movement between otherwise isolated populations
could substantiallyreduce their vulnerability.
Research effort should be directed at identifying where
corridors potentially exist and documenting their use(e.g., Projects 3-4 and SO).
6. Protected areas cover only a minority of cat species
ranges; most cats occur outside of protected areas, inhuman-modified habitat. Conservation of cats is thus
inextricably linked to development in both rich andpoor countries. Cat specialists should become more
involved in the development process. Their role is to
study conflicts between people and cats, and to identify ways to reduce those conflicts so that both people
and predators can coexist. This is the subject of thefollowing chapter.

and Conclusions

1. Most cats are associated with forest habitat. Tropicalmoist and dry forests are particularly important. Morevulnerable (Categories l-3) species are associated withtropical rain forest than any other habitat type. Tropical
Asia has undergone the highest rate of deforestation in
the 1980s and its moist and dry forests harbor more
vulnerable species (Categories l-3) than any other179

Part II: Major

Issues.

Chapter

2. Managementof Big Cats Near

PeoplePart II
Major Issues in Cat Conservation

Management

Chapter 2of Big Cats Near People
economic liability, people take the law into their own
hands. This chapter thus focuses on two strategies toencourage people to tolerate the presence of cats.
First, the problem of cat predation, on both livestock
and people, is reviewed, and a number of measures which
can be taken to mitigate predation are put forward.
Management to reduce the impact of predation on livestock is one of the most important elements of a cat conservation strategy. The extent of the predation problem isreviewed, and a number of mitigating measures evaluated.
Such measures are aimed at halting direct persecution ofcats, on the assumption that, if predation problems are few,
local people and land owners will leave cats alone.
However, if livestock is safely protected, and there are noalternate suitable wild prey available, cats will still not beable to persist in such areas.
It will not be possible for most governments to set asidem uch more of their wild lands from development. The
second strategy, th.erefore, aims to provi de incentives forlandow ners to mai ntain wi ld lands with cat populations,rather than convert land to forms of use which are inhospitable to cats. Since the conversion of wild lands is taking
place for reasons which are primarily economic, so too
must be the incentives to conserve them instead. This is an
emerging field in wildlife conservation, and pilot projects
have been established based upon local people earningmoney from the sustainable use of species ranging fromtrees to iguanas to butterflies to antelope. However, economic incentives to specifically conserve cats are few,
since cats more often have costs than value. Two of themain options (tourism and hunting) are reviewed, and further application discussed. A third, commercial trade incat products, is reviewed in Chapter 4.

Introduction'

Conflict between livestock owners and predators has
existed since food animals were first domesticated around
9,000 years ago. Domesticated animals have reduced
escape abilities compared to wild herbivores, and are
exceptionally vulnerable to predation, which is a problem
wherever wild cats and livestock share range. The previous chapter showed that most cats occur outside protectedareas, and that most habitat outside protected areas has
been modified by humans. Livestock husbandry is a major
form of land use, and if livestock replaces wild prey it
becomes the chief suitable sustenance for cats. This isespecially true for big cats-many smaller cats subsist on
rodents, which can increase under some forms of agriculture and livestock husbandry. Peoples’ persecution of thelarger cats for predation on livestock, or for the danger
they can pose to human life, is the final step in the processof their disappearance outside protected areas, which starts
with habitat loss and fragmentation. To prevent further
erosion of cat range and numbers, ways must be found forpeople and cats to live together.
Laws against indiscriminate killing of big cats are oneof the primary reasons that populations still exist near people. Enforcement of protective measures for cats will
always be a fundamental component of managing these
populations, and conservation of cats living near people
could be greatly improved with sufficient resources andtraining for national and local government wildlife authorities. This is particularly true for developing countries
with impoverished people and limited resources to spend
on wildlife conservation. It is imperative that countries
which have well-managed and successful wildlife protection programs increase their financial and technical assistance to those which do not. This can only be of benefit
cat conservation.
However, laws protecting cats and their prey fromindiscriminate killing are often not sufficient to curb thefundamental pressures leading to their decline outside protected areas. Where cats are perceived as a nuisance or an

Cat Predationtoon Livestock

Persecution by humans in response to livestock predation,
both actual and potential, has been a major factor resulting in the disappearance of big cats from large areas of180

Par? II: Major

their former range in historical times, including the puma
from eastern North America, the tiger from most of China,
and the lion from north Africa and southwest Asia. In
many cases, the big cats were declared vermin and bounties paid by the government for skins. The recent, rapid
global increase in human numbers, and the concomitant
spread of settlement, agriculture and other activities, havewidened the arena where conflict between cats and people occurs. In India, for example, which supports 15% of
the world’s cattle and over half the world’s tigers, thehuman population has risen by nearly 50% since the initiation of Project Tiger in 1973 (Jackson 1993a).
Most cats-all the big cats, and many of the smalltake livestock, from horses and cattle to poultry.
Especially with regard to the big cats, livestock predators
are found to have been crippled by injuries, old age, or disease, which could have been a handicap to their taking
wild prey. Jim Corbett (1944), the famous hunter of tigers
and leopards turned man-eaters in India, was one of thefirst to point out the link. Rabinowitz (1986) in Belizeand Hoogesteijn et al. (1993) in Venezuela reported that
more than half the jaguars killed for livestock predationthat they had examined had old head or body wounds.
Most of the problem jaguars (n=l3) examined by
Rabinowitz (1986) had head wounds, but none of theskulls of 17 non-problem jaguars had injuries. However,healthy cats also take livestock (Hopkins 1989).
Some studies have shown that stock-raiding cats are
more likely to be males than females (Suminski 1982,

Cattle on ranches in the Venezuelan
by ranchers as a result.

llanosare attacked

by jaguars

Issues.

Chapter

2. Managementof Big Cats Near

PeopleRabinowitz 1986), and more likely to be sub-adult thanadult (Rabinowitz 1986). Others, such as Bown’s (1985)
survey of North American management authorities on
puma predation, have found that adults of both sexes are
more often implicated than younger animals. Perhaps
Sitton’s (1977) observation that it is impossible to predict
which cats will turn to stock raiding is still valid (Lindzey1987). Both Suminski (1982) and Rabinowitz (1986)
observed that some big cats ignored available livestock
prey, while others became “addicted.” Of nine leopards
monitored by Mizutani (1993) living on a Kenya ranch(where wild ungulate prey was available), only one habitually took livestock.
On the other hand, in the special circumstances ofranching on the outskirts of protected areas, the most likely
stock raiders are dispersing sub-adults. For example,
Anderson (198 1) reported that male lions were destroyedoutside the borders of South Africa’s HluhluweKJmfolozi
Game Reserve complex twice as often as females. Most of
the males were less than three years old (20 out of 25).
Sub-adult males, having left the reserve, seldom returned,
whereas there were over 50 incidents of females with cubsleaving and returning immediately. Males were foundmuch further from the park boundary (mean of 23 km)
than females (mean of 0.6 km). Stander (1990) classifiedlions from Namibia’s Etosha National Park which took
livestock on border ranches as either occasional raiders(lions with known histories and no previous record ofstock raiding) or problem animals (habitual raiders). The

and pumas,

181

which

are persecutedPart II: Major

Issues.

Chapter

2. Managementof Big Cats Near

Peopleet al. 1994). Ginsberg and MacDonald (1990) came 40 a
similar conclusion in their review of livestock predation by
canids. Estimates of annual stock losses to pumas in the
United States range from a high of 7% of sheep in a problem area of southwestern Utah (Bowns 1985) to a low of
0.3% of sheep in Nevada (Suminski 1982). Evans (1983)
concluded that verified puma predations affect fewer than
1% of ranches in New Mexico on an annual basis.
Similarly, predation by African wildcat and caracal
accounted for the loss of only 1.1% (1,508 animals) of
small livestock held by farmers in the Nuwe-Roggeveld
district of South Africa’s Cape province in 1984- 1985
(Vorster 1987). In Namibia, interviews with ranchers by
the Veterinary Services Department of the Ministry ofAgriculture (unpubl. data, Cheetah Conservation Fund)show that leopards and cheetahs were each responsible
for an average annual loss of approximately 320 cattle and375 sheep and goats between 1986 and 199 1. On anational level, the yearly losses to these two cats combined
is equivalent to only 0.03% of total cattle holdings, and
0.01% of total small stock.
However, the impact of cat predation is highly variable on the scale of individual livestock operations. For
example, Hoogesteijn et al. (1993) compared cattle deaths
on Venezuelan ranches. On the first ranch, deaths due tobig cat predation were considerably fewer than othercauses of calf mortality, and accounted for only 6% of all
losses or deaths. The ranch was well-managed but, following the owner’s declaration of a ban on hunting jaguar,
the proportion of deaths attributable to cats rose to 15%

sex and age structure of occasional raiders was similar tothat of the entire Etosha population, with slightly more
sub-adults of both sexes. Most problem animals, on theother hand, were sub-adult and adult males. Stander(1990: 41) described these lions as “very wild and difficult to immobilize or destroy. They were...apparently
aware of the dangers involved.” In India, sub-adult lions(both male and female) were significantly more likely thanadults to raid livestock on the outskirts of the Gir Lion
Sanctuary. Saberwal et al. (1994) concluded that the Girlion population is at saturation density, and the sub-adults
were seeking both food and territory.
Cat species differ in their livestock prey and mode of
capture. For example, on a ranch in Brazil, pumas killed
mainly calves, with some sheep, while jaguar kills were
33% calves, 57% cows, and 10% oxen and bulls(Crawshaw and Quigley in prep.). Mizutani (1993) lookedat predation by lions, leopards, and cheetahs on a mixed
livestock ranch in the Kenyan highlands. One leopard
climbed into the newborn calf enclosure to take calves, on
average one per month. Leopards also killed stray animals
left outside the fenced enclosures at night. Lions roared
outside the fenced enclosures holding cows and steers,causing the animals to panic and sometimes break throughthe fence. Cheetahs took sheep rather than cattle, attacking
during the day when they were spread out grazing. All
three cats together took about 1% of the ranch’s total stockon an annual basis.
On a large scale, livestock losses to cats are generally
low, of the order of <I -3% of total stock per year (Jackson

Goats,
forest,nicknamed“Billy the desert-maker,”in a Nepalese
which is the vital habitat of tigers and their prey.

182

wildlife

reserveprevent

regenerationof thePart II: Major

Issues.

Chapter

2. Managementof Big Cats Near

Peopleanimals (Sitton et al. 1978, Bowns 1985, Jalkotzy et al.
1992). In South Africa, Stuart (1986) found that 73% of
104 livestock kills by leopards and 22% of 79 caracal kills(mostly sheep) in Cape province were of two or more
stock animals. One leopard killed 5 1 sheep and lambs,
while caracals have been recorded to kill 21 and 22 sheepin single incidents (Skinner 1979, Stuart 1986). Other
spectacular examples include single pumas killing 59
sheep (mostly lambs) in Nevada (Suminski 1982), and 420
chickens in Canada (Jalkotzy et al. 1992). In Tibet, thekilling of 107 sheep by one snow leopard in a single night
was reported (Jackson et al. 1994), and a snow leopard in
Ladakh was known to have killed 40 sheep and goats (Foxand Chundawat 1988). Guggisberg (1961) reports six
lions killing 51 ostriches kept in an enclosure. In all theseincidents, only a small proportion of the animals killedwere actually eaten.
To sum up, livestock predation is a significant problem on a local, rather than national or regional, level. It
causes the greatest amount of economic hardship in
poorer, developing regions with few livestock per household. Predation incidents can arouse considerable hostility
toward wild cats.

(about 40 calves per year). However, the owner maintained the hunting ban. On another ranch, smaller than
the first and situated in a more agriculturally developedarea, cat predation accounted for 3 1% of calf losses ordeaths (between l-6 calves per year). Big cats were resident on the first ranch, but not on the second.
While the economic impact of livestock predation canbe significant even for larger industrialized ranches, on asmaller scale, particularly in developing countries, the lossof just a few domestic animals can be a major setback.
Oli (1994) surveyed villages in Nepal’s Manang District,situated within the Annapurna Conservation Area, thelargest reserve in Nepal for snow leopards. He recorded 72
animals lost to snow leopards in 1989- 1990, representing
2.6% of the livestock held by 102 households, with a total
value of U.S. $3,866. This represents an average household loss of 0.7 animals, valued at about $38-a substantial amount for the local people in a country where theaverage rural annual income is just $122 (Anon. 1990a),among the lowest in the world. Some households wereaffected more than others. One family lost two adulthorses in one attack, with an economic value equivalent
to six times the average rural annual income, while anotherlost 11 goats, representing 20% of their livestock. These
losses were serious enough to upset the household
economies for several years. The high value of theselosses led to considerable antipathy toward snow leopardson the part of local villagers: 97% wished to see snow
leopards eradicated.
Similarly, the area of Namibia formerly known as
Bushmanland is largely undeveloped, with only subsistance livestock husbandry. In eastern Bushmanland, there
are only 29 villages, with average stock holdings of 16 cattle and two horses. In 1992-1993, lions killed livestockon nine occasions, taking eight cattle (1.7% of the total
number kept in eastern Bushmanland) and four horses(9.3% of the total). The affected villages suffered an economic loss of U.S. $56 (Stander 1993). However, theJu/‘hoan residents have little cash income, and still depend
mainly on hunting (20%) and gathering (80%). The loss
chiefly represents a setback to efforts to establish a sustainable cattle industry for food supplementation (Anon.
1992d). The losses contributed to the desire of 84% of villagers surveyed that lions be eliminated (Stander 1993).
Local antipathy toward cats as predators is magnified
by the phenomenon of surplus killing. When a cat breaks
into a fenced enclosure and encounters large numbers of
prey animals, it will sometimes kill more than it needs toeat. Fenced animals cannot escape the predator, and their
panicked movements repeatedly stimulate the cat’s killing
instincts. Surplus killing is a rather frequent phenomenon; it is common for pumas in North America to kill morethan one animal per attack if small stock (sheep, goats,
poultry) are involved, but with cattle, most kills are single

Management to Minimize the
Problem of Livestock Losses
to Big Cats
The traditional response to livestock predation has been
to attempt to eliminate all predators in the area. Nowak( 1976) has documented that government-sponsored predator control efforts in western North America were responsible for the death of nearly 67,000 pumas between 1907
and 1978. However, it is rather surprising, given historical
success in eradicating populations, that more modern
attempts have generally failed (Lindzey 1987). For example, the complete elimination of pumas from problem
regions in New Mexico has been attempted three timestwice to protect domestic sheep and once to protect wild
sheep. None of these removals resulted in a reduction of
predation (Evans 1983), and pumas are still there today.
One reason may be that the vacuum was rapidly filled byimmigrants from neighboring areas, such as occurred
when lions were culled from a section of South Africa’s
Kruger National Park (Smuts 1978). Another may be that
big cats eventually adapt to persecution, and become more
secretive and more difficult to catch.
Nowadays, management measures to minimize livestock predation take three forms: attempts to eliminate thespecific animals causing the damage; improved anti-predator and general livestock management; and compensation
for livestock lost to predators.

183

Part II: Major

Problem

Chapter

2. Management

AnimalControl

Issues.

of Big Cats Near

Peoplecurrently listed as protected game, and land owners may
legally kill them in response to livestock predation withno official reporting requirement.
Both approaches have their problems. The problem in
Namibia is that the management strategy does not prevent
indiscriminate killing, resulting in the loss of large numbers of cheetahs which are not problem animals. Most
cheetahs end up being shot in a trap, rather than killed in
the act of predation (L. Marker-Kraus in Zitt. 1993). Over5,600 cheetahs were killed in the period 1980- 199 1,
according to official statistics (Govt. of Namibia 1992).
Some ranchers have gone to a lot of trouble: over the same
period, one rancher killed 175 cheetahs on his property,
an average of 16 animals a year (Kraus and Marker-Kraus
1992). The number of cheetahs killed annually in Namibia
shows a declining trend, down from 850 in 1982 to 301 in
1990, probably indicating a declining population.
In India, it can take a long time for an official to arriveat the scene of the predation, and in practice they often
never arrive. The degree to which people take the law
into their own hands and kill big cats is unknown. Becauseof the danger of increasing illegal persecution of cats,
Johnsingh et al. (1991) and Saberwal et al. ( 1994) urgethat the authorities attempt to eliminate problem animals,
as quickly as possible, irrespective of the species involved.
In practice, problem animal control is most efficient,in terms of minimizing livestock predation as well as conserving cats, when the actual problem animal is targetted.
It is best to target problem animals specifically because
they may lead others to kill livestock. For example, with

In most cat range states, it is permissible for predators
which take livestock to be killed or removed. Regulations
vary as to whether the livestock owner himself may takeaction, or must call upon a government animal control officer to do the killing, and to what degree predation must
be verified before elimination of the problem animal issanctioned. The following examples give an indication of
the range of variation.
In India, predation on cattle is common news in all partsof the country (Sawarkar 1986), which supports 15% of
the world’s people, 15% of the world’s cattle, and 54% of
the world’s tigers. The government has prohibited thekilling of big cats: the only exception is elimination of aconfirmed, deliberate man-eater. In extreme (and rare)cases of persistent livestock predation, the offending animal may be captured and either translocated to a reserve or
given to a zoo (S. Deb Roy, pers.comm.).
In Namibia, cheetahs are significant problem animals,
and land owners are permitted to kill cheetahs “to protect
the life of livestock, poultry, or domestic animal...while thelife of such livestock is actually being threatened.” Theowner is then required to report to the nearest wildlife
authority within 10 days to obtain a permit for possessionof the skin (Nature Conservation Ordinance, 1975). The
government does not verify specific predation incidents,
although predation is monitored in a general way throughtwice-annual visits of veterinary control officers to every
commercial ranch in Namibia. Lions and leopards are not

Puma captured

for attacking

livestock184

on a Venezuelan

ranch.

Part II: Major

Box 1Predators

Issues.

Chapter

2. Managementof Big Cats Near

Peopleand Farmers

South Africa’s Endangered Wildlife Trust has produced
an information booklet for livestock keepers entitledPredators and Farmers (Bowland
et al. 1993). The
booklet is useful and the information is well presented:
it deserves imitation elsewhere. The first part consists ofbrief accounts, with distribution
maps and photos, oflarge and small carnivores found in the region, with
notes on their beneficiai activities. The booklet explains
the importance of determining whether the prey animalrNas killed by a predator or died of other causes and wasiust fed upon, and explains how to determine this byskinning the carcass and looking for mortal wounds andother evidence.

predators, cat species are much more likely to feed
upon a prey animal’s inner and/or outer thighs in a“clean and tidy” manner. Paw prints and hairs of eachcarnivore species are drawn ciearly to aid diagnosis.
The book emphasizes that predator control efforts canbe costly, and that before undertaking
any action a
farmer should compare the costs of losses to predators with the costs of predator control. He may find hecan live with certain levels of predation, but more oftensome form of predator control is necessary.
Predators
and Farmers
is available
from theEndangered
Wildlife Trust, Private Bag x 11, Parkview
2122, South Africa.

The book contains photos illustrating the various predator “signatures”:
for example, in comparison
to otherlions, Stander (1990) found that occasionalraiders sometimes accompaniedproblem males on forays into cattle
ranches. Femaleswhich take to livestock predation may
teachtheir cubsto kill stock.
While it is best to have an expert tracker and identify
the animal, this is not always feasible. Many cats will
return to a kill if it is left alone: this is probably the best
way to identify the actual killer, although the kill may beinvestigated by another, innocent predator. Well-trained
dogsmay alsobe able to pick up and follow the scenttrail
of a cat which doesnot return to a kill (Stuart 1982).
One innovation for targeting problem animals is the
toxic collar, a broad collar put aroundthe necksof domestic animals, with a capsuleof powerful poison attachedwhere the collar runs under the animal’sthroat. Cats generally kill larger livestock with a throat bite, and are poisonedwhen they bite through the collar and its capsule.
Such collarsareusedin somedevelopedcountries,including France, the U.S., and South Africa. This allows the
livestock owner to target actual livestock killers, without
having to involve a governmentproblem animal officersuchpeople,if they exist at all, are usually too few or too
busy to respond to all reported predation incidents efficiently. Collars could be made available to ranchers atlow, subsidizedrates.
A similar techniqueis the placing of traps, either lethal
or steel-jawtraps, nearlivestock areas,but thesetraps areoften indiscriminate and may catch innocent animals. F.
Mizutani (in Zitt. 1993) is experimenting with aversion
training by injecting the carcassesof livestock killed byleopardswith the nauseatingsubstancelithium chloride.
One leopard which killed a sheepcame back to eat thetreatedcarcassand did not return to livestock killing.
Rather than being eliminated, problem animalscan betranslocated.But translocationshave met with mixed success (Hamilton 1976, Seidensticker et al. 1976, Mills
1991, Anderson 1992). Habitual problem animalsoften
return to stock-killing (Rabinowitz 1986, Stander 1990).
Stander (1990) was able to return occasional raiders totheir homerangeswithin EtoshaNational Park, with only
one of 12 translocated lions resuming stock raiding.
However, in practice, the original home range of a wandering predator will seldombe known. Translocation is
discussedin greaterdetail in Chapter 6.

Improving
Livestock

General Anti-predator
Management

Livestock owners may exaggeratetheir lossesto big catsby including lossesdue to other causes,including poor
management techniques. Even on the conservationminded ranch in Venezuela mentioned earlier, where
jaguar hunting was banned despite an increase in calf
losses,ranch managersestimatedannuallossesto big catsat 200 animals,but careful investigation of carcassesby
Hoogesteijnet al. (1993) showedthat only 40 calvesa year
were killed by cats. Hoogesteijnet al. (1993) listed othercausesof calf lossas:injuries inflicted during round-up ortransportation, snakebite, drowning, disease,and slaugh-

185

Pat? II: Major

Issues.

Chapter

2. Managementof Big Cats Near

Peopleter for human consumption. Mizutani (1993) adds theft
to the list. L. Marker-Kraus (in Zitt. 1993) cites leg injuriescaused by falling into burrows (common in arid regions),
abortions in cattle and small stock, poisonous plants, andbirthing problems with first-time heifers. Other predators
may turn out to have a more significant impact than cats: ina problem zone in southwestern Utah, for example, it wasestimated that coyotes were responsible for 92% of sheep
losses, and pumas for 7% (Bowns 1985). In California,
domestic dogs are responsible for more sheep deaths thanpumas (Trulio 1989).
By improving basic livestock management, owners can
reduce losses to causes other than big cats, and increase
their profitability. Such measures include vaccination of
livestock against disease, and improved husbandry measures to increase pregnancy and juvenile survival rates.
For example, Hoogesteijn et al. (1993) noted that only 4050% of beef cattle pregnancies in the Venezuelan llanos
are successful, and only 30-40% of calves reach breedingage. L. Marker Kraus (in Zitt. 1993) reports a 55% calving rate in Namibia; in other words, 45% of heifers that
could reproduce do not do so.
With specific regard to improving anti-predator management, the following general principles can be recommended (Stuart 1982, Evans 1983, Bowns 1985, Ginsberg
and MacDonald 1990, Oli 199 1, Kraus and Marker-Klaus
1992, Quigley and Crawshaw 1992, Bowland et al. 1983,
Hoogesteijn et al. 1993, Mizutani 1993, Jackson et al.
1994; see Box):
0

0

0

0

0

0

0

0

0

0

0

Mountain

Proper disposal of livestock carcasses so that predators
do not acquire a taste for livestock;
Changing from cow-calf to steer operations where
losses to big cats are heavy;
Guards or guard dogs for daytime grazing (or even, as
the Cheetah Conservation Fund of Namibia has suggested, donkeys or baboons);
Controlling birth seasons rather than allowing births totake place randomly;
Keeping cows and calves under closer supervision
when calves are young, and away from areas of thickvegetation or rough terrain where cats may lurk;
Keeping, rather than selling or trading, experienced
herd lead animals, so that they can teach appropriately
cautious behavior to younger animals;
Keeping a few cows or steers with horns in the calving
herd;
Rounding up livestock at night into soundly fenced
enclosures and posting armed guards with lights;
Improving the security of fenced enclosures throughbetter fencing, including, where economic, electric;
Permitting wild prey species to co-exist with domestic
livestock;
Fencing off ranch areas which adjoin prime cat habitat, or avoiding grazing in such area.

lion killed in Utah.

Compensation

for Livestock

LossesPaying compensation for livestock losses is a way of
encouraging land owners or local people to tolerate predator presence. In some places, compensation is paid by thenational or local government, while in others, compensation is paid by conservation groups. Compensation canbe an effective tool when it is not abused. Several compensation progams are reviewed below.
When lynx were reintroduced in France, Switzerland,and Austria, sheep losses provoked great hostility fromfarmers towards lynx conservation. Although sheep losses
were low (400 lost in Switzerland over 15 years) compared
to annual livestock losses to lynx and other predators elsewhere in Europe, farmers raised a great outcry which
threatened the success of lynx reintroduction. Compensation was paid voluntarily in Switzerland by the Swiss
League for Protection of Nature, and in France by WWFFrance. In 1988, the national and local Swiss governments
took responsibility for paying compensation. At the present time, Switzerland pays out about U.S. $18,000 annually as compensation for lynx kills, a small sum comparedwith the nearly $35,000,000 invested annually as a subsidy
for sheep husbandry. Compensation is thus not a major
economic outlay, and has helped to smooth public accep186

Part II: Major

tance of the lynx (Breitenmoser
and BreitenmoserWiirsten 1990).
In North America, two American states (Wyoming andColorado) and one Canadian province (Alberta) pay compensation for livestock losses to pumas. In Alberta, theLivestock Predator Compensation Program covers only
food-producing stock (i.e. cases of predation on dogs orhorses are not covered), and market value of a loss must
exceed CA $100 (U.S. $72) per calendar year. Reported
incidents of predation are investigated by provincial
wildlife officials, and claims for compensation are
reviewed by one of two regional committees composed ofprivate producers and government representatives fromanimal health, production, and wildlife management interests. Losses are judged as “confirmed kill,” “probablekill,” or “missing animal,” and include livestock fatality,
injury from which recovery is deemed improbable, and thedisappearance of animals in conjunction with confirmed
kills or injuries. Since June 1990, the program has paid
100% compensation for confirmed kills, 50% compensation for probable kills, and no compensation for missing
animals (previously, 30% compensation was paid). From
1974- 1987, 6 1 claims for puma predation were approvedfor compensation, an average of 4.4 per year. For every
approved puma claim, there were five wolf, 13 bear, and42 coyote claims approved between 1974- 1983 (Pall etal. 1988). Annually, total compensation for puma kills
averages only CA $1,617 (U.S. $1,171: Jalkotzy et al.
1992). In Colorado, annual compensation paid for puma
kills is considerably higher: U.S. $45,000 in 1990-1991(Hansen 1992). Hansen (1992: 62) interviewed J. Talbott,
Assistant Chief Warden for the Wyoming Fish and Game
Department, about the effectiveness of compensation interms of puma conservation.
Talbott believed that
Wyoming’s program “increases tolerance of the cougars
and helps ranchers view them as a bona fide wildlife
resource that people like to see out there.”
A major issue in North America is that puma predation often takes place on minimally supervised public
lands leased to private individuals for livestock grazing.
Expenses of livestock compensation programs extend
considerably beyond the actual compensation paid out,
and include costs of monitoring, verification, negotiationof compensation, and administration. In its puma management plan, the government of Alberta province(Jalkotzy et al. 1992) gives predator control measures lowest priority on public grazing leases, although compensation is still paid. The California-based
Mountain LionFoundation suggests that occasional losses of livestock topumas on grazing lands leased by the public be tolerated
by the owners as “part of the price of doing business”(Hansen 1992: 111).
In India, state governments are responsible for dealing
with livestock predation according to guidelines issued by

Issues.

Chapter

2. Managementof Big Cats Near

Peoplethe central government. When an incident is reported, awildlife official carries out an investigation in the presenceof the complainant and some other villagers. No compensation is payable if the incident occurred inside a protected
area, or the carcass has been disturbed in any way. The
amount paid out varies from state to state and is related tothe age and sex of the animal and its intrinsic value-e.g.,compensation is higher for milch cows and buffaloes, anddraught animals. It is difficult to evaluate the success ofthese programs. In the Gir Forest area, Joslin (1984)
declared that the compensation scheme for livestock losses
was of limited practical benefit to herdsmen. A decade
later, the situation had not improved: 81% of Gir area villagers interviewed by Saberwal et al. (1994) said they did
not bother to file claims. The problems they cited included
low level of compensation in comparison to the purchaseprice of replacement animals; excessive travel to reportlosses; the likelihood that an official would not be available
to register a report of livestock predation within the mandatory reporting period (24 hours); subjective assessments
by officials of the worth of predated livestock; and difficulties associated with receiving payment for settlements.
Somewhat similar to official compensation is private
insurance for livestock losses. However, it is doubtful
whether any individual owner sustains high enough losses
from cat predation to warrant paying for such insurance.
On the other hand, in at least one area of Canada’s
province of British Columbia, sheep losses to coyote predation were so high that insurance coverage was withdrawn (I. Ross in litt. 1993).
Paying livestock compensation can be a relatively lowcost way to encourage livestock owners to tolerate thepresence of cats, for, even with the most advanced stock
husbandry, some losses to predators are likely to occur.
Oli (1991) reviewed various management options available to reduce livestock predation by snow leopards in
Nepal’s Annapurna Conservation Area, and concluded
that a livestock compensation fund, locally administered,
had the best potential to reduce the conflict between local
people and snow leopards. He suggested that a Snow
Leopard Conservation Committee be formed of prominentvillage representatives, an NGO representative (Annapurna Conservation Area Project), and the local wildlife
authority. It would be up to the Committee to develop thedetails of the scheme (procedures for claim, levels of reimbursement, etc.) and inform the villagers about it. Oli suggested that compensation be paid only for verified snow
leopard livestock kills in cases where appropriate measures
were taken to avoid predators, and not be paid for the lossof male yaks or oxen left unattended throughout the year,
the loss of an animal not corralled at night, or the loss of anold, injured, diseased, or otherwise dying animal. The
scheme should be simple enough to permit illiterate villagers to make claims properly, but effective enough to

187

Part II: Major

Issues.

Chapter

2. Managementof Big Cats Near

Peopledetect and discourage abuse.
Oli discussed both the pros and cons of his compensation idea, which are a useful summary for compensation
schemes in general. The merits of the compensation program include: (1) it is the management method mostacceptable to the community (as snow leopards cannot
legally be eradicated); (2) it involves a direct financial
incentive, so that livestock owners are more likely to cooperate with laws protecting the snow leopard; (3) the
scheme involves an endowment fund, with the interest
used to pay compensation, so that it is sustainable; (4) the
management committee will consist mainly of local representatives, so conservationists will not solely be held
responsible for any perceived shortcomings in the scheme;(5) as the management committee will act as a liaison
between the conservation authority and local people, it willaid implementation of other conservation measures.
Drawbacks of compensation include: ( 1) livestock
losses from any cause may have to be compensatedbecause it is impossible to go to the site and determine theactual cause of death on all occasions; (2) false claims
could be difficult to detect, and compensation of such
would set a bad precedent; (3) it is possible that local people will accept compensation but continue to kill snow
leopards secretly, and it would be difficult to determinethat this was happening; (4) management committee members might use their position to political advantage, leading
to a general loss of faith in the compensation scheme; (5) if
the committee fails to function efficiently and impartially,
it will reflect badly on the conservation authority.

Touristmeets

a tiger in India.

Programs Which Make
Wild Lands an EconomicallyCompetitive
Form ofLand Use
Conservation of cats outside protected areas will be a battle of attrition, especially for the big cats, where the wild
prey base has been eliminated. If wild ungulates are killed
off to make room for cattle, there is little hope that cat populations will persist in these areas. If protective livestock
management works, there will be nothing for the cats to eat.
For this reason, the recent and increasing interest on thepart of the conservation community in promoting ways inwhich wild lands, with their full complement of species,
can be an economically competitive form of land use, isof great significance. Perhaps the best example of how
successful such policies can be are the state-owned andprivate hunting concessions in North America and Europe.
Considerable revenue is generated from the sale of hunting
licenses and other associated infrastructure for the budgets of wildlife authorities. Tourism is also significant, but
not to such a high degree as in protected areas.
In terms of cat conservation, however, the most promising examples are projects in developing countries which
work with local people so that use of wild lands is not only
sustainable, but also profitable. Such programs concentrate on the economic and resource values of wild plants
and animals, particularly the large ungulates. A key con-Controlled

188

tourism

can promotewildlife

conservation.

Par? II: Major

Tourists viewing lions on a zebra kill in east Africa,
exchange,
but requires stricter control.

wherewildlife

tourism

is a major

Issues.

sourceChapter

2. Managementof Big Cats Near

Peopleof foreign

demand as a result of conservation campaigns. The only
examples of managed commercial cat fur harvest, the bobcat and Canadian lynx, are discussed in Chapter 4. The
only other economic values of cats are tourism and trophy
hunting: these are discussed below, with specific reference
to their contribution toward conservation of cheetahs onprivate land in Namibia as a case study.

cept is that landowners must benefit economically fromwildlife if they are to refrain from converting wild habitat
to other forms of land use. This concept is operative not
only in development-oriented programs like Zimbabwe’s
CAMPFIRE(Communal Areas Management Plan for
Indigenous Resources), but also in a growing commercial
interest in converting African cattle farms to game farms.
While these new developments in conservation are
exciting, their contribution to cat conservation has yet to be
proven. Cats themselves have limited economic value,
and most game farms and organized wildlife producer
communities rely on the value of large wild ungulates asmeat and trophy animals, although there are numerous
other values to be derived from wild animals and plants.
While it is these types of schemes which have the greatest
potential to contribute to cat conservation outside protected areas (since populations of cat prey species are conserved), it is possible that local managers will see cat
predation much the way livestock owners do: as an unacceptable cost of doing business. In southern Africa, forexample, most game farms, especially those managed formeat production, but also those with valuable trophy antelope, do not tolerate big cats.
Specific economic incentives to conserve cats are few.
One is the value of their fur. This value is limited byinternational trade restrictions
and declining public

Tourismand Trophy

Hunting

There are few places in the world, apart from Savannah andsome other non-forested parts of sub-Saharan Africa,
where cats are easily visible, although the knowledge thatthey are present, and the chance of catching a fleeting
glimpse and finding tracks, adds a special attraction towildlife areas. Where tourists can expect to see big cats,these species have a significant role, and generate revenue.
Over a decade ago, Thresher (1982) calculated the tourist
value of a male lion in Kenya’s Amboseli National Park
to be U.S. $128,750. He surveyed tourists visiting thepark, found that most desired to see full-grown male lions
with “proper” manes, and that they spent about 2.5% of
their game-watchingtime observing such lions. He
derived his dollar figure by allocating responsibility for
2.5% of the park’s annual income to male lions, divided by

189

Part II: Major

Issues.

Chapter

2. Managementof Big Cats Near

Peopletioned, game farm managers are often opposed to keeping
cheetahs and other predators because they prey on valuable
antelope, usually purchased by the manager.
Tourism is an industry subject to a host of vagaries,
most importantly the state of the economy of developed
countries from where most tourists originate. While
tourism is currently on the rise in Namibia, future growthis unpredictable, and in the end has little to do with cheetahs. It can be concluded from this discussion that tourism
does make a contribution to conservation of cheetahs onprivate lands, but it is a small one. Moreover, the morecountries with cheetahs on private lands which duplicate
Namibia’s private game farm strategy, the more likely thevalue of a cheetah will fall due to competition for a limited resource (tourists).
A number of African countries permit sport hunting oflion, leopard, and cheetah (Martin and de Meulenaer 1988).
For cheetah and leopard, which are on CITES Appendix I,
hunters are able to take their trophies back home under aquota system approved by CITES. Trophy hunters spend
considerably more per capita in host countries than dotourists: Makombe (1994) notes that the price tag for a 21day lion hunting safari in Tanzania is around U.S. $35,000,
much more than a tourist would spend visiting the country. Moreover, hunting revenues can make a substantial
contribution to governmental income. For example, in
1990, government revenues from hunting licenses in
Tanzania amounted to some U.S. $4,500,000, more than
twice the revenue earned from the national parks system($1,900,000: Makombe 1994). Revenue from sport hunting can be a key part of sustainable use of wildlands
schemes. In a review of sport hunting as a sustainable useof wildlife, Edwards and Allen (1992) report that communities participating in the (CAMPFIRE)
program earned$4,000,000 from sport hunting in 1990- 199 1, representing
about $400 per household. In some communities, this represented a doubling of annual household income.
Namibia has a well-developed sport hunting industry.
From 1983- 199 1, safari hunters took an average of 2 1
cheetahs per year (Govt. of Namibia 1992). Most of thesewere shot on private land, although mainly on game farms
or hunting concessions rather than commercial cattle
farms or communal land. While the total amount ofmoney spent by a trophy hunter during his stay can beconsiderable, the trophy price for a cheetah is currently
rather low, with a high of U.S. $1,500 and a low of $600.
In contrast, zoos sell live cheetahs for prices between U.S.
$6,000-10,000. Moreover, in some cases, trophy animals
were purchased from farmers who had trapped them,rather than tracked and hunted on a game farm (L. Marker
Kraus in litt. 1993).
In order for trophy hunting to play anything but a minor
role in cheetah conservation, cheetah trophy hunting
would have to expand a good deal, especially into cattle

the number of lions in the park, and multiplied by a lifespan of 10 years. This is, of course, a highly theoretical
exercise. The park would not necessarily lose 2.5% of its
income if lions were to decline or disappear-althoughitmight lose more.
Martin and de Meulenaer (1988) calculated that a leopard was worth U.S. $50,000 annually to the privatelyowned Londolozi Game Reserve in South Africa,
assuming the highly-visible leopard there to be responsible
for one-fifth of the net profit of U.S. $250,000 a year.
However, they declare that most countries in Africa losemoney from wildlife tourism when the full costs are taken
into account, thus suggesting that leopards in general havenegative value.
In India, the secretive tiger has become visible in a few
reserves as a result of effective protection and the habituation of some individuals to the approach of people invehicles or on elephants. These reserves have generated
tourist revenue but, apart from some employment in hotels
or as drivers, local communities have not benefitted, andthe wildlife authorities have not received any significant
contribution towards reserve management costs.
Namibia is home to an estimated 2,500 cheetahs, withthe vast majority of the population found on private commercial cattle ranches (Morsbach 1987). Such private
lands, where other large predators (lion and hyena) havedisappeared, are thought to be of critical importance toconserving viable populations of cheetah, whose numbers
may be limited by these competing predators in reserves.
Wildlife tourism (including both camera and rifle safaris)
is a major sector of the Namibian economy (Berry 1991c),
and a recently established NGO, the Cheetah Conservation
Fund, is attempting to promote the country as “the cheetah capital of the world.”
A major tenet of national
wildlife policy is that landowners are encouraged to maintain wildlife on their property and to reap economic benefits from it. Cheetahs in Namibia are thus an ideal casestudy of the contribution economic value can make to cat
conservation. If such a strategy is to work anywhere in
the world, Namibia stands a good chance.
However, tourism on private land faces stiff competition from larger, more spectacular protected areas, and catson private game farms or on relatively undeveloped communal lands almost assuredly attract fewer tourists than inprotected areas. If tourists coming to Namibia want to see
cheetahs and can make only one stop, they are far more
likely to go to Etosha National Park rather than a gamefarm. The decision to visit a game farm is likely to be
more related to the quality of amenities offered rather thanthe presence of cheetahs. Many guest farms in Namibia
keep semi-tame cheetahs around their premises. Such animals are a strong attraction after the hands-off policies ofnational parks and reserves, but game farms can keep tamecheetahs without keeping wild ones. As previously men-190

Pat? II: Major

ranches where most cheetahs occur and cause the mostproblems. The Cheetah Conservation Fund has been holding discussions with the Professional Hunters Associationof Namibia regarding increasing promotion of the cheetah as a trophy animal along with raising the trophy fee,with a portion of the fee to be channeled back into cheetah conservation (L. Marker-Kraus, pers. comm. 1994).
While both tourism and trophy hunting help conserve
cheetahs on private land in Namibia, their contribution atpresent is of limited utility, and does not begin to compensate for the hundreds of cheetahs killed annually on
cattle ranches as problem animals. If the contribution of
economic use options for the conservation of cats outside
protected areas is to be increased, specific cooperative
effort on the part of conservationists and the tourism andtrophy hunting industries is required.
Martin and de Meulenaer (1988) stress that sport hunting should not be considered a replacement for problem
animal control in Africa because livestock predation tends
to occur at times when hunters are not active, and they do
not particularly like to hunt in heavily-settled rural areas.
On the other hand, Swank and Teer (1988) proposed limited and controlled sport hunting of problem jaguars in
Belize, Brazil, Mexico, and Venezuela, arguing that thefees could provide a major portion of the finances neededby government agencies to implement management measures and, in part, to compensate local people for cooperating in the plan. In India, Saberwal et al. (1994)
considered the problem of local antipathy to problem
Asiatic lions in the Gir Forest to be so serious that they alsosuggested that the sport hunting option be considered,despite the fact that the subspecies is very rare, and thatno sport hunting of big cats is permitted in India. None of
the countries named has implemented these proposals.

Summary

Issues.

Chapter

2. Managementof Big Cats Near

Peoplestock management, and paying compensation for livestock taken by cats. There is relatively little material
available about how to implement these measureseffectively (e.g., see Action Plan Projects 5, 32,52,74,
93, and 102 in Part III).
4. Big cats are also persecuted because they are a danger
to people, and in some cases tigers, lions, and leopards
have become man-eaters, specializing in human prey.
In general, however, most attacks on people are theresult of accidental, rather than deliberate, confrontations. A number of suggestions are made for how people can avoid or escape big cat attacks (see Box 1, Big
Cat Attacks on People).
5. Even if livestock predation can be minimized, thechances of cats surviving on unprotected lands are slim
if there are no wild prey there to support them. For thisreason, the growing movement to develop economic
incentives to make maintenance of wild lands a viable
alternative to agriculture and stock husbandry is a welcome development. In most cases, conservation of wild
ungulate populations for meat or trophy hunting is oneof the best incentives, and this is of great potential significance for cat conservation. However, managers ofwild game are likely to be as hostile toward cats as
managers of domestic livestock, viewing predation as
an unnecessary cost. Cat specialists should become
involved in such cases to test ways to encourage theconservation of cats under these circumstances (seeAction Plan Project 22).
6. The economic values represented by cats themselves
are few, consisting of tourism and hunting. Commercial hunting in North America for the fur trade isreviewed in Chapter 4. Tourism and trophy hunting
can be an incentive for the conservation of cats on
unprotected lands, but their impact is limited at present.
If tourism and trophy hunting are to play a major role incat conservation, it will require that cat specialists cooperate with the industries to work toward this goal (seeAction Plan Projects 31 and 33).

and Conclusions

1. Most of the world’s cats occur outside of protectedareas, where the most direct threat is human persecution. For the larger cats especially, active management
and protection measures are needed to ensure their conservation in areas used by people.

7. It is difficult to develop recommendations on conservation of cats outside protected areas, in part because
there is no professional disclipine which embraces the
subject. Conservation of viable big cat populations
near people is highly complex but also extremely
important. The Cat Specialist Group has recognized
this (CSG 1984, McNeely 1991), and a number ofgroup members work on resolving human-predator
conflicts, but none full-time. There needs to be more
recognition of the significance of this subject, anddevelopment of a professional capacity to address the
problem (see Action Plan Project 6).

2. Predation of livestock is the primary cause of people’s
intolerance of cats. Although on a large scale predation
rates are fairly insignificant, averaging O. l-3%, theirimportance is magnified at the level of individual owners, especially in developing countries. To poor people,
the loss of just a few animals represents a major financial setback, and provides a strong incentive for extermination of cat populations.
3. Measures to minimize livestock predation include problem anim al control, improving general anti-predator
191

Pad II: Major

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Chapter

2. Managementof Big Cats Near

PeopleBox 2
Big Cat Attacks on People
The Nature and Extent of the Problem

occurs, and has been documented
by Chakrabarty(1992). Hendrichs (I 975) and Chakrabarty (I 992) speculate that the aggressive nature of the Sundarbans tigers
could be due to the effects of excessive salt imbibedwhile drinking. But this is discounted by others, and the
Sundarbans
tigers’ historical predilection
for humans
could perhaps be more simply attributed to local “culture,” with cubs learning to identify humans as prey fromtheir mothers.

Tigers, lions, and leopards have long been notorious
for fatal attacks on people. They are frequently termed“man-eaters,” but the term is, in general, unjustified. The
existence of dedicated man-eaters is a fact, but manyfatal attacks are the result of a cat’s defensive reaction to
the surprise appearance of a human.
“Tiger” and “man-eater” have become almost synonymous, although the number of attacks is remarkably low
when the opportunity
offered by human presence istaken into account. One of the worst affected areas,
the Sundarbans mangrove forests in eastern India, has
an unofficial death toll of about 100 people per yearalthough this is perhaps exaggerated,the official average being 36. While high, these figures have to be set
against the fact that 35,000 people move through the
Sundarbans each year (Chakrabarty 1992).

Given existing patterns of human use of the Sundarbans
forests and the peculiar habitat conditions that exist,
man-eating is unlikely to be eliminated, although it has
declined to some extent in recent years as a result ofbetter regulation of human activities and special measures to deter tigers. However, as already noted, tigers
attack a relatively small percentage of the large numberof people who knowingly venture into dangerous areasof the Sundarbans,and there are no records of tigers
going out of the forests in search of human prey.

McDougal(1987) points out that there are certain
regions where conflict between people and tigers has
been historically minimal, and cases of man-eating arecomparatively
rare. He mentions Burma, Malaysia,
Sumatra, Thailand, and Vietnam. On the other hand,southern and northeastern China, Singapore, and India
have had serious and persistent man-eater problems.

The second case involves tigers moving into settled andcultivated areas. Dudhwa National Park, at the foot of
the Himalayasin Lakhimpur-Kheri
district of Uttar
Pradesh, has become increasingly isolated from previously contiguousareas of wild habitat in the last 40
years. Surrounding grasslands have been converted tosugarcane cultivation right to the edge of the park. The
park is home to a healthy tiger population, but dispersal
to other areas has been disrupted by intensive humansettlement and activity. Sugarcane fields attract wild
ungulates from the park, which are followed by tigers.
Since the typical cultivation
regime of sugarcane
involves long periods of tranquility and relatively little
human activity in the fields, tigers tend to become resident and breed there, preying on any wild ungulates orcattle in the area. In such a situation, there is a high possibility of confrontations
between tigers and people
entering the fields. Between 1978-l 988, 197 people
were killed and 33 “man-eating”
tigers shot in the
Dudhwa area (Khushwah 1990).

At present, two areas are noted for tiger attacks: theSundarbans,and the district of Kheri in northern India,
bordering southwestern Nepal. The two areas illustratedifferent aspects of the problem.
The Sundarbans
is an example of people moving into
tiger range. There is no permanent habitation in theinterior of the Sundarbans.
People enter the forest to
fish, cut wood, and to gather honey and other forest
products. The only drinking water available is saline;
high tides submerge large areas twice daily; and protruding spiked air-breathingroots make walking difficult. Nevertheless,although the area appears to hold
ample wild prey, Sundarbans tigers have for centuries
been known for unusually aggressive attacks on humanvisitors and deliberate man-eating.
A French traveller
in India in the 17th century, Franqois Bernier, recounted
how tigers took fisherman from their boats (Bernier1670). In 1791, a British hunter was taken by a tiger inbroad daylight while lunchingwith friends(The
Gentleman’s
Magazine
1793).
Such behavior still

Lions in both Africa and India have attacked people andbecome man-eaters.
Despite an epidemic of attacks inthe early years of the 20th century, the Asian lions of
the Gir Forest in Saurashtra,
western India, havebecome renowned for their passivity. However, a newwave of attacks began in 1988, and up to March 1991
Continued on next page

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Part II: Major

lions had mauled 120 people, of whom 20 died (Ravi
Chellam and Johnsingh 199313). The attacks have been
linked to the decline of the local cattle population during
a prolonged drought. The surviving cattle have been
more carefully protected. This has led to more aggressive hunting by the lions, for which livestock has long
formed an important part of diet. They have broken into
the compounds of houses where livestock is kept at nightby tearing off roof tiles. The lions have thus come into
conflict with people, resulting in injuries and death. In areview of the problem, Saberwal et al. (I 990) noted thatthe 1901-1904 epidemic of lion attacks (during which atleast 66 people were killed) also followed a drought.

Issues.

Chapter

2. Managementof Big Cats Near

ungulates, so that lions deprived of natural prey turned
to livestock and entered villages, where they killed people (Anon. 1987b).
E. Marshall Thomas (1990, 1994), whose family has along history of association with Bushman groups living in
the Kalahari Desert, offers an interesting insight into whylions may be peaceable or hostile toward humans. In
the 196Os, when she lived there, and for thousands ofyears previously, the lions and the Bushman people had
an “arrangement.”
They hunted the same prey andmade use of the same waterholes,
both a precious
resource in the Kalahari. She points out that the average group size of Ju/hoan people and the average pride
size of lions were of equivalent weight, so that a meat
meal sufficient for a group of people would also satisfy
a group of lions. The lions and the people occasionallyhelped themselves to each other’s kills, and compromised regarding the waterholes:
the people occupied
them during the day, and the lions during the night. No
one could remember an incident of a lion attacking aperson, with the exception of a paraplegic girl who was
killed, and Thomas knew of no lion killed by a Bushman,

Conflict between lions and people in Africa has been lesswell documented, probably because news media are notso developed as in India. Guggisberg (1961) has documented a number of attacks on people throughout thecontinent, and the infamous man-eatersof Tsavo in
Kenya (Patterson 1907) still vie for public attention withthe man-eating tigers of Kumaon in India (Corbett 1944).
During the 198Os, fatal attacks by lions in southern
Tanzania
were attributed to heavy poachingof wild

Continued on next page

Nepalese villager ki lled in an accidental
in Kathmandu
Zoo.

confrontation

Peoplewith a tiger, which was later captured

193

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Part II: Major

Issues.

Chapter

2. Managementof Big Cats Near

PeopleWild cheetahs have also never been reported to attack
people, although recently a cheetah which got loose ina zoo in Mississippi attacked a child, who escaped serious injury by “playing
dead” (P. Jackson,pers.
comm.).ln
North America, puma attacks on people,
while unusual, have increased in recent years. Beier(1991) documented
53 unprovoked attacks in the U.S.
and Canada between 1980 and 1990, of which nine
were fatal. They included the killing of a five-year-old
boy in 1989 and an 18-year-old jogger in 1991. Thirtyof the 53 attacks occurred in British Columbia, with 20
on Vancouver
Island. Hornocker(I 992) speculates
that long-term intensive hunting of pumas on the island
has selected
the most aggressive
as survivors,
although retaliation is usually thought to lead to cats
becoming more cautious. The attacks appear to haveincreased
as suburban
settlementsin the west
encroach on puma habitat (Foreman 1992, Seidensticker and Lumpkin 1992).

although everyone had heard of lions attacking Bantu
people, who kept cattle and hunted lions. Thomas
writes, “No one can explain the truce, because no one
understands it. The truce was simply taken for granted,
as most situations involving animals are simply taken for
granted. Animals are assumed to be static in nature.
So even today, with both the human and the animal populations stressed and damaged, few people realize thedifference between how things are now and how things
once were.”
For comparison,she describes the unpredictableandsometimes
aggressive
behavior of lions in Etosha
National Park, where Hei//kum Bushman people had
lived until the 1960s. Her explanation is that the lions
there lost their cultural heritage of how to live amicablywith humans, once the people who lived there were
removed, and the people who came afterwards(theThe first
tourists) were confined to automobiles.
recorded incident of “tourist’‘-eating
recently took placein Etosha (Nowell ef al. 1994).

Measures
The leopard

is a naturally secretive big cat, which
is able to live in surprisingly close proximity to humans,
and leopards which turn to man-eating
have proved
extremely difficult to hunt down (Corbett 1948, TurnbullKemp 1967). Attacks tend to be nocturnal.
In India,
leopards have claimed victims in rooms full of sleeping
people without causing any disturbance (Corbett 1944).
Other people have fallen victim to lurking man-eating
leopards when leaving their houses to relieve themselves at night. Leopard attacks on people have always
been common in hill areas of India and Nepal, whererugged terrain provides cover. They have also occurred
around dwellings in the vicinity of the Gir Forest (Ravi
Chellam in lift. 1992).

to Minimize

Risks to People

While there are no guaranteed ways to avoid attacks by
big cats on humans, the following suggestions may be
helpful:*There is little documentationof jaguar attacks onpeople, but A. Rabinowitz (in litt.) reports hearing manystories in Belize, Brazil, and Guatemala that lead him
to believe that, under the right circumstances,
attacks
could happen. Forest dwelling Indians in Latin America
say that jaguars take children (A. Michelangeli,
pers.
comm.).
Snow leopards have no record of killing people. Womenand unarmed villagers, including teenagers, may chasesnow leopards away from the goats they have killed in
Himalayan villages without fear of attacks, while cornered snow leopards are surprisingly subdued towards
humans throwing stones at them (Mallon 1987, R.
Jackson, pers. comm.).

Where big cats are liable to threaten people, circumspect behavior is called for. People in groups are
generally safe, but children especially excite the interest of big cats and are more vulnerable (65% of 58
known victims of puma attacks were less than 16
years old: Beier 1993). Care should be taken where
cover may conceal a resting cat. A warning shout orother noise to announce
an approach
is likely tocause it to move away. For example, many people
living in wild environments
are purposefully
noisy
when walking from place to place in order to scareoff potentially dangerous animals. Talking quietly toor yelling at a threatening
cat may scare it off.
Thomas (1990) describes the way Ju/hoan Bushmen
spoke to lions on a kill which they intended to rob,
speaking “firmly but respectfully.”
Beier (1993:409)
describes how a 50.year-old female hiker, attacked
and knocked down by a puma, set up her backpack
as a shield, faced the puma, and (in her words)“began talking to her the way you would if you were
trying to soothe a dog or cat.” She kept this up for 30
minutes until she heard other hikers nearby andyelled for help; their noisy approach (with one hiker
blowing a whistle) scared the cat away.

Continued

194

on next pagePart II: Major

0

0

l

l

Issues.

Chapter

2. Managementof Big Cats Near

PeopleWhen threatened by a big cat, it is dangerous to turnand run-in at least two cases, running appeared tostimulate a puma to select the victim out of a larger
group.
However, running up a tree might help,
although in one puma attack, the two cats climbed
after the girl, who kicked them and hit them with astick to make them leave (Beier 1993).
Seidensticker and Lumpkin (1992) stress the importance of maintaining eye contact with a threatening
cat, as well as taking care not to bend or squat. tn the
Sundarbans,the use of face masks, worn on theback of the head, has proved a deterrent to tigers,
who usually attack from behind. Large eye spots on
the back of a cap may also be effective.
Throwing objects at an attacking cat may help and,once attacked, 20 of 29 puma victims successfully
fought off the cats with bare hands, a stick, a knife, ajacket, or a rock. In several cases, even children
unassisted by adults were able to repel pumas byfighting back. In several other cases, children carriedoff by pumas obeyed their parents’ instructions to“play dead” and were limp. The pumas, although carrying the children, did not attempt to kill them, andpeople were able to scare them off (Beier 1993).
Attacks should be investigated immediately to establish whether they were accidental (perhaps arisingfrom the cat being surprised), or a case of deliberate
hunting. If an attack is clearly determined to be deliberate, the wisest course of action is to eliminate theproblem animal as soon as possible.
If the attack
appears to be the result of a surprise confrontation, no
action need be taken, unless the body has been eaten.

Woodsmanin the Indian Sundarbans
back of his head to deter man-eating

Big cat attacks are perhaps most likely to occur in settled areas bordering protected populations of cats. A
healthy population will include a dispersing class ofboth sub-adult animals seeking to establish a territory,
6 and older former residents which have lost theirs.
Both of these types of cats are prime candidates forbecoming
problem animals: the sub-adults
have
immature hunting skills, and the hunting efficiency ofolder animals is declining. If risks to people (or livestock) are unacceptably high in such situations, there
may be no solution but a program of eliminationofthese “surplus” cats, which is generally what occurs,in an illegal, informal and indiscriminate
fashion. In
Nepal, for example, tigers and leopards which strayout of the Royal Chitwan National Park are often poisoned by livestock owners (Seidensticker et al. 1991,
C. McDougal pers. comm.).
l

195

wearing
tigers.

a mask on thePart II: Major

Issues.

Chapfer

3. ResearchPart II
Major Issues in Cat Conservation

Chapter 3
Research
Introduction

species, 63%). The priority species for which research
effort has been high or very high are all big cats (cheetah,
tiger, lion, snow leopard), which have both economic
value in terms of tourism and trophy hunting, and cost,when they come into conflict with human interests, primarily livestock farming. In fact, all species for which
research effort has been high or very high are “revenue
earners” with legal value (nine species) and/or “revenue
consumers” with problem animal status (seven species).
Most species of conservation concern (14 species, 73% ofCategory l-3 species) have no legal economic value.
Research effort has thus not been altogether strategic
regarding conservation of the Felidae, being strongly correlated with legal economic value rather than species vulnerability. This is not necessarily a conscious choice byresearchers. The big cats have been relatively well studied
primarily because of their position at the top of ecologicalfood chains, observability, and charisma. However, it isalso these qualities which give them legal value (throughtourist and trophy hunter interest) and cost (through predation of large livestock or man-eating). In addition, at firstglance, the small spotted cats exploited for their fur would
appear to be exceptions, as they have been commerciallyhunted in large numbers while there has been virtually noresearch effort. However, for the Latin American cats,the major period of exploitation took place lo-20 years ago(see Chapter 4, Trade), and was largely illegal. National
government authorities, which often fund research, did notbenefit. In the case of the leopard cat, which is legally harvested, the Chinese government has recently recognized
the need for sustainable management, and has substantiallyreduced harvest and export quotas while a major research
project is organized (Johnson and Fuller 1992, Johnson etal. 1993).
The conservation benefits of linking the ecologicalvalue of catsto an economic value have beenemphasizedin Chapter2. Many, if not all, of the specieswhich are ofconservation concern have the potential to help meet thecostsof their conservation through development of revenue-earning options, tourism being currently the mostMany priority projects in this Action Plan (Part III) fallunder the aegis of research, although the reason for thismay not be immediately obvious in a document concerned
foremost with species conservation. To some, a call for“more studies” is seen as an obstacle to real conservation
which, for cats (as discussed in Chapters 1 and 2), primarily involves maintaining a network of protected areas andimproving conditions in the human-modified areas whichlie in between. However, knowledge of basic natural history, including diet and habitat requirements, is scanty formost species, let alone subspecies. This will impede
attempts to determine suitability of habitat corridors, andto prevent extirpation of cats from areas used by people. In
the words of Caughley (1994), it is important that conservationists have confidence that their background knowledge of species’ natural history is “adequate to avoid silly
mistakes.”
Reliable gauges of animal abundance areimportant for planning conservation actions, and forassessing their actual impact. Long-term studies are necessary to provide meaningful insight into species biology.
Finally, while the science of conservation biology-incorporating computer models which simulate population
dynamics; molecular research; identification of uniquely
adapted sub-populations; and evaluation of the effects of
disease upon populations -continues to advance throughthe 1990s the lack of data for most cat species means they
will be left out unless field research efforts are intensified.
Table 1 summarizes the last few decades of researchon cats, including field, captive, and laboratory studies.
In general, research effort has been low (low or very low= 20 species, 56% of the 36 species in the family Felidae).
Research effort has been exceptionally high and thoroughfor the northern group of species, the lynxes. Intraspecificbiological and situational variation has been poorly represented for most cats (24 species; 67%). Most importantly,
research effort has not been strategically focused. Effort
has been low or very low for those species of priority conservation concern (vulnerability
rankings l-3; n= 12

196

Part II: Major

Issues.

Chapter

3. Research

Table 1
Research Effort for Cat Species
Species

RarityRanking

Research
Effort’

Legal
Value2Distribution3

Representationof Variability4

Cheetah, A. jubatus
Caracal, C. caracal
Bornean bay cat, C. badia
Asiatic golden cat, C. temmincki
Chinese mountain cat, F. bieti
Jungle cat, F. chaus
Sand cat, F. margarita
Black-footed cat, F. nigripes
Wildcat, F. silvestris
Jaguarundi, H. yaguarondi
Ocelot, L. pardalis
Oncilla, L. tigrirws
Margay, L, wiedi
Serval, L. serval
Canada lynx, L. canadensis
Eurasian lynx, L. lynx
lberian lynx, L. pardinus
Bobcat, 1. rufus
Pampas cat, 0. coloco~oGeoffroy’s cat, 0. geoffroyi
Kodkod, 0. guigna
Andean mountain cat, 0. jacobitus
Manul, 0. manul
Leopard cat, P. bengalensis
Flat-headed cat, P, planiceps
Rusty-spotted cat, P. rubiginosus
Fishing cat, P, viverrinus
African golden cat, P. aurata
Puma, P. concolor
Clouded leopard, N. nebulosa
Lion, P. leo
Jaguar, P. onca
Leopard, P. pardus
Tiger, P. tigris
Marbled cat, P. marmorata
Snow leopard, U. uncia

3(A)
5b
2
3
2
5b
4
2
5c
5c
5a
3
4
4
4
5b
I
5a
5a
4
2
2
4
5b
2
3
2
2
WA)
3(A)
3(A)
WV
WV
2(A)
3
WV

High
Int.
v. low
v. low
v. low
Low
Low
v. low
Int.
v. low
Low
v. low
v. low
Low
High
High
Int.
V. highv. low
Low
v. low
v. low
v. low
Low
v. low
v. low
v. low
v. low
V. highLow
V. highlnt.
HighV. highv. low
V. high-L tf, P
P
0
0
0
0
0
0
P
P
0
0
0
0
F
f, p
0
F, H
0
0
0
0
F
F
0
0
0
0
J-9 l-4 p
0
J-7 H, p
T H, pT H, P
T p
0-f-f H, p

s N
s, N
T
-L E
E
T E, Ns, N
Ss, N E
A
A
A
A
s, N
A
E
E
A
A
A
A
A
E
-L E
T
T
T, E
S
A
T E
S
A
s, N T
T
-L E
E

Poor
Poor
Poor
Poor
Poor
Poor
Poor
Poor
Int.
Poor
Good
Poor
Poor
Poor
Good
Int.
Good
Good
Poor
Poor
Poor
Poor
Poor
Poor
Poor
Poor
Poor
Poor
Int.
Poor
Int.
Int.
Int.
Int.
Poor
Good

1 The felid bibliography compiled by Foreman et al, (1988) is largely based on sources from North America, and* contains many studies on captive animals in addition to articles from the popular press, but serves as a usefulindex to the relative degree of attention paid per species. In several cases, additional studies not included in
the bibliography are accounted for. Very high = 2OU+ publications; High = t OO-200; Intermediate = 50-100;
Low = 15-50; Very low = cl 5.
2 F = Fur trade; T = Tourism; H = Trophy hunting; P = Problem animal. This column is an index of the importance ofa species to national governments by way of its revenue-earning potential (or vice versa, in the case of species
considered problem animals). Tourism revenue is attributed to a species only when it is likely that the majority
of tourists visiting protected areas would be likely to specifically desire and expect to see that species,
3 S=Sub-Saharan Africa; N = North Africa and Southwest Asia; E = Eurasia; T = Tropical Asia; A = The Americas.
Q An indication of how well a species’ diversity of habitat types or situation (e.g., inside vs. outside protected areas)
is represented by studies conducted. lnt. = Intermediate.

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Part II: Major

Issues.

Chapter

3. Researchfeasible and significant. Here research needs to play amajor role. With appropriate advertising, tourists can be
encouraged to visit reserves, public or private, that contain a rare and little-known (to the public) species of cat
which they might see. However, cats are primarily nocturnal, and small cats are notoriously elusive and difficult
to observe, particularly in densely vegetated environments(such as tropical rainforests, which are currently experiencing a boom in tourist interest). Without knowing more
about the habits of these species, it will be impossible tocreate situations in which tourists would be likely to get a
glimpse (and a photo) of these animals, let alone a
background lecture on their ecology. Increasing tourist
interest in the small cats would provide a tangible demonstration of these species’ value to the governmental authorities charged with their conservation.
Potential economic value is of course not the only
aspect of wild cats which requires further study. This
chapter is concerned with those areas of scientific researchwhich are important for cat conservation. Progress to dateis reviewed, and areas where improvement is needed arehighlighted. Both field and laboratory studies are covered.
The study of captive animals is discussed in Chapter 5.

Table 2
Species for Which Fewer than Three
Adequate Studies of Natural History
Have Been Done
Species

Sub-Saharan Africa

38%

Black-footed cat, wildcat,
African golden cat

North Africa and SouthwestAsia

100%

Cheetah, caracal, jungle cat,sand cat, wildcat, setval, leopard
33%

Eurasia
Chinese mountaincat, manul

TropicalAsia

82%

Bornean bay cat, Asiatic golden cat,jungle cat, leopard cat, flat-headed
cat, rusty-spotted cat, fishing cat,clouded leopard, marbled cat

Field Studies
Field studies needed for cat species are discussed in four
categories below, in order of increasing complexity. For
all categories, the conservation importance of studying
cat populations in settled or otherwise human-disturbed
areas is emphasized. Progress to date is reviewed for twoof the research topics, but data collected from natural history studies are given in the Species Accounts, and studies of human/carnivore conflicts are covered in Chapter 2.

NaturalPercentage ofTotal Number ofSpecies Occurring

TheAmericas

58%

Jaguarundi, oncilla, margay,
pampas cat, Geoff roy’s cat,kodkod, Andean mountain cat

Latin America. Study is lacking for those specieswhichlive in areasremote from urban civilization, such ashighmountains,deserts,or tropical rain forest.
Why hassciencepassedby somany of the cat species?
In part, it is becausecatsare relatively difficult to studythey have evolved, in terms of both morphology andbehavior, to avoid detection. Studying the ecology of anocturnal cat would be practically impossiblewithout theaid of radiotelemetry,andthis technology hasonly become
efficient andreliablesincethe 1980s.Also, in order to collar a cat, it mustfirst be caught, andthe literature of the Cat
SpecialistGroup is replete with examplesof the difficultyof live-trapping cats for study purposes.
Someindividuals arenotoriously trap-shy, while others
are caught repeatedly. For example, Vaughan (in press)
describesan attempt to mount a study of the ecology ofthree small cats in Costa Rica’s La Selva Biological
Station. Using chickens, mice, fish, and meat as bait,

History

Natural history represents the fundamental definition of
species uniqueness in ecological terms. Natural history
involves study of where (distribution, habitat selectivity) and how (diet, activity patterns, social organization)a species lives. Natural history studies do not always
lend themselves to testing hypotheses, and for this reason
have been very much neglected by the increasingly specialized students and scientists from the academic
research community.
As shown in Table 2, a basic understanding of the biology and ecology of most of the small cats is lacking. Thisis especially true for the more vulnerable cats. However,even the common species have not been studied. Regionally, natural history research is most needed for the catsof north Africa and southwest Asia, tropical Asia, and198

Part II: Major

Issues.

Chapter

3. Researchresearchers ran a seven km trapline with 25 box traps, but
after four months succeeded only in catching opossums
and ants. On the other hand, J. Beltran caught four ocelots
in one day in southwestern Texas, where the species is
actually quite rare (Anon. 199 1c). One of five snow leopards radio-collared in western Nepal was caught five times(Jackson and Hillard 1986). While Rabinowitz (1992)
caught the same leopard several times in western Thailand,
Jenny (1993) had enormous problems catching just one in
the Ivory Coast’s Tai’ National Park. Problems in capture
translate to problems in obtaining an adequate and balanced (in terms of sex and age) study sample.
Other ways in which cats do not lend themselves toconvenient study include the length of time often required
to gather sufficient data, when there is inevitably a variety
of deadlines to meet which bear little relationship to therhythms of the study population. Cats can be wide-ranging, and this complicates logistics. Cats are nocturnal,
while humans are not. Also, even if radio-collared, study
subjects are usually rarely, if ever, seen by the researcher.
This is not only frustrating, but also potentially limits thedata which can be collected.
However, all these setbacks apply equally to the northem hemisphere cats -the lynxes, the puma, the European
wildcat-allof which have been relatively well-studied.
Research on these cats has been largely funded by national
governments, and it is safe to conclude that the main reason why most cats in other regions have not been studied islack of funds. Most developing nations cannot allocate
sufficient funds for basic research, and international conservation groups are less likely now than ever before tofund this type of work, preferring instead to support studies aimed at “biodiversity” rather than single species.
These obstacles are not insurmountable. Regarding thefunding problem, this Action Plan is designed to highlightpriorities within the cat world for both seekers of projects
and donors of funds (Part III). With regard to the wider
conservation benefits of research as well as to the difficulties of studying cats, attention should be drawn to theimportance of including local people as project participants. Research needs are greatest in the developing countries. Involving local people as active participants in astudy and providing training in the basics of field observation makes for a valuable transfer of technology and expertise to less developed areas. Conversely, local knowledgeand “bush skills,” which exist among traditional hunters
and trappers can make a significant, but often overlooked,
contribution to the success of a studyand may turnpoachers into gamekeepers. With budgets for field studies being typically small, the hiring of local people canincrease the size of a research team for a relatively low
cost. Moreover, such action can be invaluable for shaping
local perceptions regarding the value of their environment
and the cats within. This has important long-term consequences for what happens after the principal researcher
departs. If more well-trained and motivated individuals

Radio-collaring

study

a tiger in Nepal’s

ChitwanNational

Park during

a long-term

199of tiger ecology.

Part II: Major

Issues.

Chapter

3. Researchunderstanding of species distribution and habitat selection,and perhaps revise what are now rather pessimistic conceptions of species status.
For example, the flat-headed cat of tropical Asia isthought to be primarily tied to wetland environments,
where it feeds on fish and crustaceans; at least 50% of wetland sites in the region are considered to be under moderate to high degrees of threat (Scott and Poole 1989,
WCMC 1992). However, there are anecdotal reports of its
presence in oil palm plantations in Malaysia, where it isthought to prey on rodents or even domestic poultry (M.
Khan in Zitt. 199 1). Similarly, the kodkod of Argentina
and Chile is thought to be strongly tied to primary moist
temperate forest, but it may do well in plantations of North
American pine species where rodents are abundant (J.
Rottman, pers. con-m-r.in Melquist 1984). As a final example, the clouded leopard is another species traditionally
thought to be tied to a particular habitat type-in this caseprimary tropical rainforest. However, it has been sighted
in logged forest (Davies and Payne 1982, Rabinowitz et al.
1987, Santiapillai and Ashby 1988, M. Khan in litt. 1991),
and may also make use of grassland and scrub habitats(Santiapillai and Ashby 1988, Dinerstein and Mehta
1989). Moreover, a cub was recovered from a tea plantation in India (Anon. 1992j). Are these animals dying inthese habitats, or living there? Are such records isolated
instances, or are the habitat requirements of these species
more flexible than presumed? This information has
important consequences for understanding species rarityand prioritization of conservation effort.
One reason why so few studies have been conductedof cats outside protected areas relates to why research islacking on cats living in remote or inaccessible habitats.
Practically speaking, in both situations establishing aworkable research schedule can be a difficult and timeconsuming process. Outside protected areas, it is necessary to work closely with a number of people who
probably understand little of conservation (unlike park
research staff), or may even be openly hostile to cats.
Where cats are subject to persecution, they can be
expected to have developed even greater secretive behaviors, magnifying the usual difficulties of capture and monitoring. Yet it is these cats for which study is of greatest
importance. In terms of science, some aspects of behavior (i.e, activity patterns and predation) are likely to differ
substantially from those inside protected areas, and understanding these differences is the key to appreciating thescope of species adaptability and evaluating probabilityof future survival (see, for example, the cheetah Species
Account). Study results can also aid the development ofstrategies to reduce conflict between cats and people, andto manage habitat or human activity in ways that allow
maintenance of cat populations at an appropriate level.
Biotelemetry equipment is indispensable to natural his-

Monitoring
the movementsof a radio-collaredtiger in Nepal’s
Chitwan National Park in order to establish its range.

around the world were actively to monitor, report on, andpromote conservation of local cat populations, the benefits to international conservation would be enormous.
Natural history should be studied in both protected anddisturbed habitats. The vast majority of studies have been
done within protected areas, in part because results canthen be evaluated in the light of a body of data co1lected onother aspects of the ecosyste m, and theories and hypotheses can be m ore readil y tested. Howe ver, it is increasingl y recogn ized that few parks are large enough toconstitute undisturbed nature. Indeed, most parks where
research is an active and ongoing process are also actively
managed, and the two complement each other. As disturbed and unprotected habitats make up the majority of
the cats’ ranges (Table 11, Chapter 1), the need to carry out
more studies in these types of areas cannot be overstated.
It is important that researchers begin to assess the ways
in which cats adjust to different forms of habitat modification and disturbance, and to identify “common denominators” which can be used to promote conservation outside
protected areas. This type of research could broaden

200

Pat? II: Major

for radio-collaring
population.

in the Jura Mountains,

Switzerland,

Chapter

3. Researchbehavior, predictability, its personality. After a while offollowing through rain and shine, night and day, thick andthin, one begins to understand, from nothing but a disembodied signal, a great deal about the animal’s moods andbehavioral tendencies. Most species have highly characteristic or typical behavior patterns that are clear only after
2-5 days of following a given sex/age class. Males andfemales are often quite different in their behaviors.”
Most cats, most of the time, are sympatric with othermembers of the family Felidae. The largest assemblages,
or guilds, of felids occur in the tropical regions of Asia, theAmericas, and Africa. The ways in which these species
coexist, and the implications for relative abundance, havereceived comparatively little research attention. It is
intriguing that in each of these tropical guilds, there are
two species identical in jaw length, including thejaguarundi and margay in the Americas, the caracal andserval in Africa, and the Asiatic golden cat and fishing cat
in southeast Asia (Kiltie 1984, Seidensticker and Lumpkin1991). This also applies to the lion and tiger in Asia (D.
Smith in Zitt. 1993), although the two are no longer sympatric, and it is doubtful that their ranges ever over-lapped
to any significant extent. Jaw length, a measurement of
maximum gape, has been shown to be strongly correlated

tory studies of cats. L. Emmons (in Zitt. 1992) has emphasized the benefits of continuously following a collared animal (as opposed to taking periodic remote locations). She
has applied this methodology to 15-20 mammal species(including ocelots: Emmons 1988) in primary tropical rain
forest, one of the most challenging environments in whichto study cats. Continuous follows, in all habitats, yielddata on the following topics: exact circadian activity, pathway and distance moved per day, habitat use (quantitative), hunting tactics (directly or indirectly), diet andpossibly kill rate, resting sites, encounters with other radiotagged animals, and exact home range boundaries.
Sampling periods should be at least several days long toallow the best understanding of how an animal really uses
its ground, although this is time-consuming and limits thenumber of animals which can feasibly be included in astudy. It is very important that the follower attempt tominimize disturbance by remaining at an appropriate distance behind the study animal and, in dense vegetation,using trails to avoid excessive noise which could “drive”
the animal or scare away other species with which it would
otherwise interact (e.g., prey). The most important aspectof this methodology, according to Emmons, is that it yields“a feeling for the animal: what it does and is, its general

Lynx trappeda reintroduced

Issues.

during201

a studyofPart II: Major

Issues.

Chapter

3. Researchanother? The largest felid guilds occur in tropical rainforest, the habitat type undergoing the highest rate of loss, andthe small cats within these guilds are very close in size.
How do they co-exist ? Ecological separation in these felid
communities can be investigated most efficiently by having a small team carry out basic natural history studies inthe same study area at the same time, with each member
concentrating on a different species.

with modal prey size (Kiltie 1984). Although these pairsof species may exploit similar-sized prey, they select different habitats. The serval and fishing cat prefer wetlands,
while the margay appears to be the most arboreal of theAmerican cats; the three others are more generalist in theiruse of habitat. The fact that for each of these pairs ofspecies, one is patterned and the other plain, may reflect
this difference (Seidensticker and Lumpkin 199 1). The
spotted morph of the Asiatic golden cat has been mostcommonly reported from China, where the fishing cat does
not occur.
Field studies have indicated that different sized cats tend
to exploit different sized prey (Seidensticker 1976, Bertram
1982, Emmons 1987, Koehler and Hornocker
1991).
Sympatric cat species may also exploit different patchesof habitat (Seidensticker 1976, Parker et al. 1983, Emmons
1987, Koehler and Homocker 199 1, Rabinowitz 1989), or
the same habitat at different times (Bertram 1982,
Rabinowitz and Nottingham 1986). To varying degrees,
these factors also determine ecological separation wherecats are sympatric with other predators (Schaller 1972,
Johnsingh 1983, Leopold and Krausman 1986, Konecny
1989, Sunquist et al. 1989, Rabinowitz and Walker 1991,
Mills and Biggs 1993, Johnson et al. in prep.).
The differences in prey and habitat selectivity between
sympatric cat species, and possibly other predators, are apriority for study because of the rapid rate of habitat modification. To what extent will this increase interspecificcompetition, and benefit one species at the expense of

Population

Status Surveys

Status surveys vary in rigor, depending on their objective,
from confirmation of species presence to estimation ofdensity or total population size. Repeated surveys allow
monitoring of population dynamics. Cats, however, arenotoriously difficult to count (e.g., Bertram 1979).
Traditional census techniques, such as transect counts or
mark-recapture studies, tend to fail when applied to cats.
The irregular, individualized, and cryptic behavior andmovements of cats violate the basic assumption of bothmethods, which is that sighting (or resighting/recapture)is predictable. More reliable techniques need to be developed which can be standardized so that results obtainedby different observers from different areas are comparable.
An important example of a move in the right direction is
the Snow Leopard International Management System(SLIMS), a protocol for standardized field survey techniques under development at the International Snow

Russian and American
researchers
threatenedby poaching.

confer202

in the Russian

Far East, where

the Amur(Siberian)tiger isPart II: Major

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3. Research

ciently assessing the distribution of sub-populations infragmented habitat. In particular, Smith et al. (1987a)emphasized the utility of GIS, which consist of computerized data bases that perform map overlays. Maps whichshow vegetation, settlement, roads, topographic features,etc.- all physical features which affect cat distributioncan be used to identify areas likely to harbor important subpopulations. Presence or absence in these areas can then be
evaluated by confirmative surveys. For example, Van
Dyke et al. (1986b) have suggested that consistent track
searches along dirt roads by competent trackers should reliably detect presence or absence of pumas in areas of suitable habitat in the eastern United States (where they are
believed to have been eradicated by the late 1800s).

Leopard Trust (R. Jackson, pers. comm. 1993).
Techniques which have been used to survey cat populations are described below. The first type, presence orabsence surveys, is important for mapping species distribution, including the identification of sub-populations. The
second type, which yields density estimates, is necessary
for understanding population status. It is a long-term goalof the Cat Specialist Group to stimulate, conduct, collectand synthesize local status surveys into a Geographic
Information System (GIS) database to map the meta-population of each species. This would greatly improve theability of the Cat Specialist Group to focus attention andresources on threatened populations. There is a long way
to go toward achieving this goal, but it is a feasible one if
the financial support necessary to refine survey techniques
and then apply them strategically is forthcoming.

Estimating
Density
Only detailed study, lasting several years and using
biotelemetry, within a relatively small area (for big cats,
generally not more than several thousand km*; for smallcats, considerably less) is likely to yield an accurate estimate of population size. In such situations, researchers canbe reasonably confident of their ability to identify all animals resident within the area, and to distinguish transients.
All other methods and estimates are less intensive, but alsoless reliable.
Estimating the size of a larger population is generally
done by taking a density estimate as described above,
expressed in terms of the number of resident adults per unit
area, and extrapolating it over other areas of similar habitat. However, Schonewald-Cox et al. (199 1) reviewed 2 14
population censuses of carnivores, and found that surveys
over relatively large areas tended to yield lower density
estimates. They concluded that extrapolating site-specificdensities over larger areas probably leads to overestimation of population size. The reason for decreasing densitywith increasing survey scale is probably a reflection of
the patchiness of suitable or optimal habitat over larger
areas (Schonewald-Cox et al. 199 1). In addition, researchers often select study sites which support relatively highnumbers of study animals, and density estimates resulting
from small-scale surveys in such areas of optimal habitat
would yield overestimates if applied to less optimal areas.
Finally, for carnivores with large home ranges relative to
study area size, more accurate density estimates may result
if some animals are included as only proportionally resident to account for parts of their home ranges which may
lie outside the study area (Garshelis 1992).
The various techniques used to establish presence orabsence can also be used indirectly to estimate densityusually in relative terms (compared to other locations or aprevious survey) rather than absolute (a numerical estimateof population size). All have drawbacks which compromise their accuracy.
Several studies have investigated whether there is a

Presence or Absence Surveys
This type of survey seeks to confirm whether a particularspecies is present in an area and provide a rough “guesstimate” of its status-rare, common, threatened, decreasing,
etc. The usual approach is to access the knowledge ofnational and local authorities, experts, and residents(Myers 1975, 1976, Teer and Swank 1977, Eaton 1978,
Melquist 1984, Hamilton 1986a,b, Tello 1986a, b,
Rabinowitz et al. 1987, Swank and Teer 1987, Martin andde Meulenaer 1988, Rabinowitz
1988, Gros 1990,
Tischendorf 199 1, Tilson 1992a, Salter 1993). Interviews
can also be conducted more intensively in areas which
potentially harbor important sub-populations. Local people, however, may not necessarily discriminate between
cat species, particularly the smaller ones. Ideally, but especially in these cases, interviews should be combined withconfirmative survey techniques that look for sign or individual animals (e.g., Koehler 1991, Rabinowitz 1993).
Both cats and people tend to use trails, and presenceor absence can often be established on the basis of sign
found alongside them, including footprints, feces, scrapes(for some of the larger cats), and tree “scratching posts”(Stuart and Stuart 1992b). When several similar-sized
species are found in the same area, however, it is preferable to use camera photo-traps. Triggered by a tripwire,a pressure pad (which can be sensitized to a minimum
weight), or an infrared beam, this method is relatively simple, inexpensive, and requires low input and maintenance(Joslin 1988, Jackson and Hillard 1986, Stuart and Stuart
199 1, Griffiths and van Schaik 1993b). Other methods
involve active attempts to attract cats: playing tape-recordings of prey sounds (e.g., goat bleating: P. Stander, pers.
comm.) or sounds of feeding (Smuts et al. 1987), scentposts, and bait (live or dead), which can be accompanied
by either a photo or sand trap (for recording track impressions), or a hidden observer.
Judicious use of maps can be very useful for more effi-203

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A variant of the sign transect is the scent-station survey,
widely used by management agencies in the United States
to census bobcats. This survey technique is preferredbecause it provides uniformity of methodology, repeatability, and cost efficiency (e.g., papers in Blum andEscherich 1979). Scent stations consist of an attractant
scent (usually a plaster disc saturated with various synthetic fatty acids) surrounded by an appropriate tracking
substrate, such as lime (CaC03).
Stations are widely
spaced along transect lines so as to minimize multiple visits by the same individual. Several studies have monitored
population size through radiotelemetry and intensive
study, and then evaluated the ability of scent-station surveys to accurately reflect population status (Rust 1980,
Conner et al. 1983, Diefenbach 1992). Conner et al.
(1983) found a good correlation, Rust (1980) found none,
and Diefenbach (1992) suggests that only multiple surveys
can detect even relatively large changes (>30%) in highdensity populations. While he recommends that a minimum of four per year be conducted, agencies generally
survey only once per year (Johnson and Pelton 1981).
Bobcat visitation rates to scent stations tend to be low(often just 2-5% of posts visited), but Diefenbach (1992)
cautions against attempts to increase visitation because thiswill statistically increase variation in resulting estimates.
Another sign survey technique is used by the Indian
government in its periodic tiger censuses, which attempt toquantify actual population size based on track uniqueness.
Pugmark outlines are traced onto glass plates in the field,
and the assumption is that differences between individual
animals will be consistently detectable (Panwar 1979, Saleand Berkmtiller 1988). This technique has been criticizedby Karanth (1993b), who tested six experienced Indian
wildlife managers by making 33 tracings of the pugmarksof four captive tigers on two different soil substrates.
While 75% of the respondents were able to correctly identify the sex of the tiger, performance was much poorer on
establishing whether the marks were made by front, rear,and left or right paws, and worst of all on estimating thetotal number of tigers responsible for the pugmarks.
Estimates were 6, 7, 13, 23, and 24 tigers-all
overestimates, the highest by as much as 600%.
However, Sale and Berkmiiller (1988) emphasize that,with intensive surveys covering a relatively small area,reasonably accurate information can be gathered regarding
the population’s age and sex structure, spatial organization, and abundance. The tracking abilities of skilledhunters, particularly those from hunter-gatherer societies,
such as the Bushmen of southern Africa, are highly developed. Such people can, with practice, learn to recognize
the spoor of individual animals (P. Stander, pers. comm.).
Scientists, however, usually cannot, and in Utah they surgically removed a rear toe from each adult resident pumain their study area to aid tracking and population monitor-

direct relationship between population density and sign
(tracks, feces, scrapes) frequency (Stephenson 1986, Van
Dyke et al. 1986b, Ahlborn and Jackson 1988, Van Sickle
and Lindzey 1992). Transect surveys can be flown (inareas of snowfall), driven (along dirt roads) or walked(along trails), and it is hoped that counting signs along
one or both sides can provide a simple index to population abundance. These studies all included intensive radiotelemetry work to provide independent confirmation ofdensity estimates based on sign alone. Although all foundsome evidence for a relationship, in practice sign frequency is unlikely to yield more than an indication of relative abundance (e.g., high or low density population), andprobably cannot be translated to a numerical estimate of
population size.
The premise of the sign transect method is that a population at high density leaves more detectable signs than apopulation at low density, but there are numerous variables, difficult to correct for, which affect how frequently
signs are deposited. For example, with regard to track frequency, some trails are more heavily used than others
because of their physical characteristics (i.e., located alongwatercourses). Also, the same individual may repeatedly
cross a trail. On the other hand, transects may be situated
so that they do not properly sample the population.
Theoretically, an accurate count would require that transects should bisect each resident individual’s home range
but, of course, this is impossible in practice (Van Sickle
and Lindzey 1992).
Scrapes and scats are also not necessarily randomly distributed because they are an important means of intraspecific communication(Leyhausen and Wolff 1959,
Hornocker 1969). To increase the likelihood of the message reaching a conspecific, sign may be concentrated attrail intersections (Robinson and Delibes 19SS), along
travel corridors such as ridgetops (Seidensticker et al.
1973), in small patches of mutually used prime habitat(Ahlborn and Jackson 1988, Fox et al. 1991a), within contact zones between territories (Seidensticker et al. 1973,
Smith et al. 1988, Rabinowitz 1989), or at communication centers, such as the “marking trees” exploited by
Namibian ranchers to live-trap large numbers of cheetah(McVittie 1979, L. Marker-Kraus, pers. comm.). Ahlbom
and Jackson (1988) found that snow leopards increased
their marking behavior when a female went into estrus.
Similarly, Corbett (1979) showed that a resident male
European wildcat increased his rate of spray-marking tenfold when a transient male ventured into his home range,and overmarked all of the intruder’s scent marks. As
marking behavior has an important social function, it islikely that it would be increased in high density populations. If frequency counts made in such situations wereused as an index, abundance of low density populations
could be underestimated (Fox et al. 199 la).

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Figure 1. Can you identify
the three left rear tracks made by
the same puma ? None of 52 participantsat the Third Mountain
Lion Workshop
in Arizona were able to. From four pumas, thethree tracks in the right column were left by one, and the three
tracks in the left column were left by three others. Measurementsfrom these tracks were included in multiple group discriminant
analysis and all tracks were correctly grouped.
Source:Smallwoodand Fitzhugh (1993).

Figure 2. Track measurements
used to identify
nine mountain lions: A. Angle between toes; B. Heel to lead toe length;
C. Heel length; D. Heel width; E. Third toe length; F. Lead toe
length; G. Lead toe width; H. Outer toes spread; I. Midline width,a parallel line 25 mm from the baseline (see K); J. Heel lobe
width; and K. Baseline used to draw midline.
Source: Smallwood
and Fitzhugh (1993).

ing (Lindzey et al. 1992).
Recently Smallwood and Fitzhugh (1993) were able todistinguish tracks of different pumas reliably by using
multiple group discriminant analysis of standardized pugmark measurements. While qualitative identification was
seldom correct (Fig. 1), and no single pugmark measurement proved sufficient, use of up to eight different measurements allowed 100% correct identification (Fig. 2).
Individual animals can also be recognized from distinctive markings, so that camera photo-traps, if distributed
widely through an area, can be used to generate rough estimates of abundance, either through direct counts of individuals or mark-recapture
techniques. Tigers can be
identified on the basis of the stripe pattern above the eye(Schaller 1967) or on the cheek (McDougal
1977).

Leopards (Bertram 1978, Miththapala et al. 1989), snow
leopards (Hillard 1989), and cheetahs (Eaton 1970, Frame
and Frame 1981, Caro and Collins 1986, Bowland 1993)
have been identified on the basis of facial spot patterns
and, for the cheetah, tail tip band patterns, which can also
be used as an index of individual relatedness (Caro andDurrant 1991). Lions can be identified by unique patternsof whisker vibrissae spots (Pennycuick and Rudnai 1970).
Species-specificbehavior can also be exploited to
attract animals for counting or marking. Smuts et al.
(1977) dragged large carcass baits to lay down scent trails,
then staked out the bait and played tape-recorded soundsof feeding at high volume to attract lions for immobilization. They succeeded in capturing and marking 409 lions
in the space of 79 nights (with two teams working each

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there could be as few as 2,750 tigers in the country (V.
Thapar, pers. comm. 1994). Just as it is unclear to what
degree tigers have increased in India since the initiationof Project Tiger in 1973 (Karanth 1993b), it is also difficult
to say to what degree tiger populations have recently
declined. If reliable gauges of tiger abundance cannot beworked out, it will be impossible to determine whether
measures such as tiger bone trade bans are effectively
reducing tiger losses to poaching.

Long-term

Tracing

the pugmark

of a tiger during

an IndianStudiesLong-term studies have been the key to insights into felid
biology and ecology. For example, the long-term studyof lions in the Serengeti has provided the data for a longand continuing exploration of why lions are social (andwhy other felids are not) (Schaller 1972, Caroco and Wolf
1975, Bertram 1978, Bygott et nl. 1979, Packer 1986,
Clark 1987, Packer and Ruttan 1988, Caro 1989, Packer
et al. 1990). The study of a puma population in Idaho
helped to define solitary felid social organization, andshowed the importance of territoriality in limiting population size (Homocker 1969, Seidensticker et al. 1973).
Moreover, long-term studies of cats have shed illumination on issues of great relevance to their conservation.
Three puma studies in North America have explored theeffects of human regulation of puma populations. Ross
and Jalkotzy (1992) studied the dynamics of a hunted population of pumas in Alberta. Two other studies have
employed an experimental approach: the effects of sport
hunting and predator control operations are being simulated by removing animals of the target sex and age
classes from well-studied protected populations (Lindzey
et al. 1992, Sweanor and Logan 1992). The “hunted” animals are actually released elsewhere, so that reintroduction of pumas (of potential interest for eastern North
America) can also be studied (Hornocker
1992). The
well-documented history of the numerous problems facedby the small Florida panther population illustrates what is
likely to happen even to protected big cats in settled areas,
unless land-use planning takes their needs into account(Maehr 1990, Maehr et al. 1991, Logan et al. 1993,
Roelke et al. 1993).
In the tropical zone, cheetahs in the Serengeti have beenstudied for over 20 years (Schaller 1972, Eaton 1974,
Frame and Frame 1981, Caro 1994, S. Durant in prep.),
and researchers have tracked a population decline, concurrent with increases in the lion and spotted hyaena populations, that challenges traditional conservation strategies
based on protected areas. Data gathered over 15 years on
tigers in Nepal’s Chitwan National Park have made thispopulation the best case study for the difficulties of conserving a population of large, dangerous cats in an insular

census.

night). Abundance of lions has also been estimated byhaving a team of widely spaced observers listen for roars,
which can be heard at distances of 4-8 km (Rodgers 1974,
Packer 1990).
In their suggestions for future directions for research
into large carnivore ecology, Quigley and Hornocker(1992: 1093) emphasized that work to improve census
techniques must continue, despite “the nature of the beast[being] against us with these secretive animals.” Much
effort has already gone into this area of research but, as
Schonewald-Cox et ~2. (1991) pointed out, the results ofmany status surveys are contained in unpublished reports
which are often difficult to access. Better communication
between researchers working in different environments
would probably not only aid improvement and standardization of survey techniques, but also help genesis andsynthesis of ideas. To this end,’ establishing the Cat
Conservation Center of the Cat Specialist Group (see Part
III) would be a significant step forward.
The current tiger poaching crisis is a prime example of
why accurate census techniques are of fundamental importance. Official tiger censuses in India put the population at
4,334 in 1989, and 3,750 in 1993 (Nath 1994), thus suggesting that some 600 tigers may have been lost in theinterval, about 20% of the population. Yet the 1989 estimate was widely considered exaggerated (Karanth 1993b),
while unofficial analysis of the 1993 data suggests that206

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Chapter

3. Research

critical for refining harvest regulations so that commercial
trapping is not unsustainable during cyclic population lows(see discussion in Chapter 4, Trade).
It is important that long-term studies remain focused on
population dynamics. Clear understanding of how a population is organized-whichanimals are actually siring
offspring and how many over a lifetime, how far the young
disperse, how many young are surviving to breeding agecan only come about through long-term study that focuses
closely on a specific population. Chepko-Sade et al.
(1987) provide useful guidelines for field collection andpresentation of data necessary to calculate the effective
size of the study population (see below for discussion).
The results of focused, long-term studies provide a baseline against which other populations can be measured, andan index which can be used to estimate the status and viability of unstudied populations with greater reliability.
Long-term studies are needed to identify the roles
played by cats within ecosystems. It is possible that thelarge cats, in particular, may be keystone species.
Terborgh (1988) suggests that jaguar and puma predationon large seed-eating neotropical herbivores (peccary, paca,
agouti) ultimately inlluences tree species composition oftropical forests.
While studies too numerous to list here have quantified cats’ offtake of prey populations, there has been no
focused long-term research on whether cat predation con-reserve surrounded by dense human settlement (McDougal 1977, Sunquist 198 1, Smith 1984, Smith et al. 1987a,
Smith and McDougal 199 1). Unpredictable environmental events which have befallen the insular population oflions in Tanzania’s Ngorongoro Crater demonstrate thevulnerability of small isolated populations to these factors(Packer et al. 1991b).
It is not surprising that there have been only a few longterm studies of the smaller cats, and that these have focusedon the lynxes. Reintroduced populations of lynx have beenstudied in Switzerland since 1983 (Haller and Breitenmoser 1986, Breitenmoser and Haller 1987, Kaczensky
1991, Breitenmoser and Baettig 1992, Haller 1992,
Breitenmoser et al. 1993, Breitenmoser and Haller 1993).
Long-term study of the Iberian lynx, the world’s most vulnerable cat species, has made it possible for researchers to
identify the causes of population decline; to propose corrective measures, and to mount an effort to accurately map
out extant sub-populations (Delibes 1979, 1980, Beltran
et aZ. 1987, Rodriguez and Delibes 1990, 1992, Palomares
et a2. 199 1, ICONA 1992). Even though there are longobservation series data on the Canada lynx, due to strong
research interest in its close relationship with cyclic snowshoe hare populations, no study has yet focused on thesocial dynamics that occur within a single population overan entire hare cycle (Breitenmoser et al. 1993b). In termsof lynx conservation, understanding of these dynamics is

Measuring

Issues.

an Indian census.

207

Pal? II: Major

Issues.

Chapter

3. Research(Joslin 1973, Rashid and David 1992, Oli et al. in press).
Since big cats will typically feed off a large carcass forseveral days, scavenging by humans should lead to anincrease in cats’ kill rates.
Habitat fragmentation could also strongly affect theamplitude of local predator-prey dynamics. When preyspecies decline due to environmental conditions in an isolated habitat block, both immigration of new prey animals
and emigration of predators seeking better food supply canbe impeded. In Africa, fences have proved to be significant
barriers to some migratory ungulates, leading to population declines (Owen and Owen 1980, Williamson andWilliamson 1985, Whyte and Joubert 1988). Within suchconfines, and under certain environmental conditions, there
is a real risk that predation could drive key prey species tothe verge of extinction. This was the justification for thedecision to cull lions from part of Kruger National Park in
the 1970s but the effort was unsuccessful-the
population
was replaced by immigrant lions, and grew rapidly under
low density conditions (Smuts 1982).
Ecosystems are complex and dynamic. Only after 30
years of study has it been possible to elucidate the characteristics of a relatively simple predator-prey system:
wolves and moose on Lake Michigan’s Isle Royale(Peterson and Page 1988). Similarly, the cyclic declineof snowshoe hare populations in Canada is a regular phenomenon which has been the focus of a prodigious amountof research attention, yet even after decades of studies,

trols these populations (Quigley and Hornocker 1992).
Cats have traditionally been and still are viewed by many
people as decimators of game. The practice of attempting
to remove all predators from reserves continued well into
the 20th century in many countries, and today predator
control is probably the main reason why cats continue tobe eradicated outside protected areas. However, generally speaking, current thinking holds that predation alone
does not cause prey declines, but can accelerate declines
caused by other factors (e.g., poor food base, or disease)(Ginsburg and MacDonald 1990). The decline could continue to the point where an insufficient prey base reduces
the number of predators, allowing the prey population torecover and increase. Alternatively, the system could stabilize with a reduced prey base until environmental conditions are such that favor prey increase. Basically, thenature of predator-prey relationships appears to be a sort offluctuating, or dynamic, equilibrium. The degree and rateof change are highly dependent on local conditions.
Human impact can have a significant influence on the
dynamics of predator-prey relationships.
Hunting byhumans can dramatically alter the species composition of
habitat which remains otherwise relatively undisturbed.
Jorgenson and Redford (1993) showed that pumas,
jaguars, and subsistence hunters in Latin America tend to
take the same mammalian prey species. People may scavenge wild prey killed by big cats (Johnsingh 1983, Thomas1990), and cats are often chased off their livestock kills

Leopardreflector

photographedby a camera
can be seen (top right).

208

trap in the Tai’ National

Park,

Ivory

Coast.

The trigger-beam

Part II: Major

Conflicts

Chapter

3. Researchdeliberately or indirectly through habitat alteration andremoval of prey species, because cats are seen as either
being valueless or, worse, parasites. For cat conservation
to succeed, two very important goals are to make wild cat
populations more valuable to both local residents andnational authorities, and to develop solutions which minimize revenue loss to problem animals. It is urgent thatthe horizons of field research on cats be broadened beyond
ecology to include socioeconomics.

researchers are still unable to determine whether the initial decline is sparked by the hares themselves (throughtheir interaction with their food resource base) or by theirpredators (Krebs et al. 1992). It may not be possible toapply the lessons of long-terms studies from one area toanother. Thus, long-term studies of the role of cats within
ecosystems will ultimately have greatest utility for the
management authority of the study area, and are most significant for high priority protected areas holding key populations of species of conservation concern. These studies
should also include the interaction between the study population and the environment immediately outside the confines of the protected area.

Resolving

Issues.

Laboratory-based

Research

IntraspecificDiversity and Systematics:
The Question of Subspecies

with People

Conflict between cat and human interests has been discussed in detail in Chapters 1 and 2. This is a key issue incat conservation, and there is a need to greatly expand
applied research, with the explicit objective of developing
and implementing workable solutions. The larger cats,
particularly those of Categories 1-3, are of top priority forthis type of field work. The topic is multi-disclipinary, andresearch will entail much more than quantification of theamount of livestock in a predator’s diet. There are unlikely
to be simple, universal prescriptions for ways in which catsand people can coexist. However, broad themes for investigation include improvement of land use planning andlivestock management, institution of appropriate controls
where cats or their prey species are being unsustainably
hunted, and development of economic use options involving cats, including tourism and sport hunting.
It is with this final topic that study of cats reaches peakcomplexity. The resolution of human/predator conflicts
straddles the two worlds of conservation and development,
which are often, but not of necessity, in opposition. This
type of research will require the largest amount of funds,
the largest number of participants, the greatest degree ofcooperation between people of possibly disparate aims,
and thus the largest possible reserve of patience and perseverance. It is not necessarily the most scientific or themost complex.
In human-modified areas, situations
change more quickly and less predictably than in protectedareas. Trying out a possible solution without having conducted a full study to evaluate its potential impact, andthen adapting management to the results, is most appropriate for work on human/predator conflicts (Stander1993). In such situations, where no action means noprogress, there is more to gain and less to lose.
This topic returns full circle to the beginning of thischapter. Research efforts have been heavily skewed
toward those species which either generate revenue or consume it as problem animals. People destroy cats, either

Review of species status (Part I) shows that the crisis at
hand for wild cats is genetic erosion within species, rather
than loss of species. No species are at present threatenedwith imminent extinction, but populations are being extirpated. However, while there is general agreement among
felid taxonomists regarding recognition of cat species,
confusion reigns on the subject of cat subspecies (seeAppendix 1 for a listing). For example, in its action plan,
the Felid Taxon Advisory Group of the American Zoo andAquarium Association(AZA) called for taxonomic
research aimed at validation of subspecific status for 235
out of 259 taxa recognized by the group (Wildt et al.
1992a: 183).
It appears that too many subspecies have been
described on the basis of too little evidence (Anon. 1991d).
For example, while 12 subspecies of bobcat have been recognized within the United States (Samson 1979, Hall
1981), Read (198 1) and Werdelin (198 1) have pointed tothe absence of geographic barriers, and suggested thatmost of these subspecies are probably invalid. Miththapala (1992: 12), after carefully documenting the different types of imprecision in description of leopard
subspecies, notes that “almost all...were defined during thelate 19th and early 20th century. During this time, it was
not uncommon for western naturalists to travel to theircolonies in the tropics on hunting trips and scientific expeditions, bring back large collections, and have a subspecies
named after themselves.” She presents preliminary evidence to indicate that leopard subspecies should be subsumed under broad regional types separated by major
geographic barriers, making the best case for treating all
leopards of sub-Saharan Africa as a single subspecies. On
the other hand, A. Kitchener (in ht. 1993) has suggested
that the snow leopard, for which two subspecies have been
described (Stroganov 1962) but are not generally recognized (e.g., Hemmer 1972, Wildt et al. 1992a), is a prime
candidate for subspeciation due to the insular and patchy

209

Part II: Major

Issues.

Box 1
Subspecies

Chapter

3. Research

Identification

Incorporating

Molecular Genetics

by Stephen J. O’Brien
To resolve the issue of cat subspecies and make thema component of cat conservation, it is first important toagree upon the basis for species and subspecies classification. My own version based on a collaboration withProfessor Ernst Mayr follows (O’Brien and Mayr 1991).

species, ordinarily under conditions of allopatry (geographical separation).
Groups of such genetically distinguishable but still reproductively compatible races aresubspecies that together would comprise a polytypic
species.

In 1940, Mayr proposed the Biological Species Concept(BSC) that defined a species as “a group of actually or
potentially interbreeding
populations that are reproductively isolated from other such groups” (Mayr 1940,1963). Reproductive isolation, the primary componentof the BSC, refers to the heritable tendency of distinct
species to avoid gene flow or interbreeding
even when
they are brought into physical contact in nature. In clarifying this notion, Mayr (1970) noted that most species
occupy distinct ecological niches, and that this ecological distinctivenessis the keystoneof evolution.
Although various alternative species concepts and criticisms have appeared(Sokal and Crovello1970,
Cracraft 1983, Paterson 1985, McKitrick and Zink 1988,
Templeton 1989, Wiley 1990) the BSC has survived thetest of time and weathered
the assault with its major
components affirmed.

Classically, a subspecies has been defined as “a geographically defined aggregate of local populations which
differ taxonomicallyfrom other subdivisionsof thespecies” (Mayr 1940, 1963, 1970). More recently, Avise
and Ball (1990) have argued that subspecies identification should be based on genetic traits. In an attempt toprovide formal criteria for subspeciesclassification,
O’Brien and Mayr (1991) suggest that members of asubspecies share: (1) a unique geographic
range orhabitat; (2) a group of phylogenetically concordant phenotypic characters that can be described; (3) a uniquenatural history relative to other subdivisionsof thespecies. Because they are below the species level, different subspecies are reproductively compatible.
They
will normally be allopatric (i.e., the reproductive barriers
are geographic), and they will exhibit recognizable phylogenetic partitioning,
because of the time-dependentaccumulationof genetic difference in the absence ofgene flow. Most subspecies will be monophyletic; however, they may also derive from ancestral subspecies
hybridization.

A major strength of the BSC is that it reflects the occurrence in natural situations of the irreversible process of
speciation.
It emphasizes reproductive isolation as thesole discriminatorof species as whole entities, b,ut
acknowledges
the occasional production of hybrid individuals, or even hybrid zones. There are numerous
examples of stable hybrid zones that appear to be geographicallybalancedby selective disadvantagesofhybrids vs. dispersal of individuals from the contact zone
(Barton and Hewitt 1985, 1989). The distinction here isthat natural occurrences of hybrid individuals or hybrid
zones between good species do not disintegratethegenetic integrity of the species as a whole, while
hybridizations between subspecies normally do produce
gene flow and genetic mixing, Reproductive isolation innature provides an effective protective device for wellintegrated genotypes.
Importantly, the BSC acknowledges the existence of appreciable
genetic diversity
within species that is often partitioned geographically(or temporally)
by population
subdivision
into sub-O’Brien and Mayr (1991) agree with H. Hemmer (inAnon. 1991 d) that the concept of a subspecies as a static unit is outdated.
According
to O’Brien and Mayr
(1991), subspecies can: (1) go extinct; (2) exchange
genes with another subspecies
and become a new“mixed” subspecies; (3) by genetic drift, selection, subdivision, or other demographic
processes change its
genetic character over time to become one or more newsubspecies;(4) if effectively isolated, become a new
species by acquiring genetic isolating mechanisms; and(5) remain unchanged.
It is not possible to know whichsubspecies wiil become new species, but they all havethis potential.
Moreover,as the time of allopatry
increases,
the probabilityof genetic differentiation
increases, and genetic differentiation is likely to include
ecologically relevant adaptations.
The possibility that

Continued on next page210

Par? II: Major

Issues.

Chapter

3. Research

genotypes that are present in all members of one subspecies, but not in any other; (2) the presence of a polymorphic
but uniquely
derivedgenetic
charactergenotype in some, but not all, individuals
of a subspecies, but not found in any other subspecies; (3) a distinct gene frequency of one or more polymorphic genetic
characters that is distinct from allelic frequencies of thesame polymorphic characters in other subspecies; and(4) geographic
isolation but absence of recognizable
genetic differentiation.
In our view, the presence of category 1 distinction is sufficient to affirm formal subspecies classification,
while anything less (categories
2-4) reflect so recent a partition as to not support subspecies designation.

subspecies carry such adaptations, coupled with theirpotential to become new species, are two compelling
reasons for affording them protection against extinction.
We are presently revisiting subspeciationin the puma
(30 classically described subspecies) and leopards (29classically described subspecies), using a suite of molecular methodologies.
Following the recommendationof Avise and Ball (1990), we are searching for discreteconcordant genetic characters that are unique to anddiagnostic for a subspecies.
When comparing several
individuals from a classically described subspecies, wehave encountered four ranked levels of distinction: (1)existence of one or more fixed genetic characters or

natureof its high mountainhabitat.
In order to conserveintraspecific diversity, there mustfirst be agreementabout how to define a subspecies(seeBox), and then identification of where subspeciesexist.
While there have, in fact, been a number of paperswritten on felid subspeciation (e.g., Pocock 1951, Weigel
1961, Herrington 1987, to name only a few), standardized criteria have not been applied. Once the identification criteria for subspecieshave beenagreedupon, efforts
to define subspeciesand map their current geographic
range should be prioritized according to either global orregional speciesvulnerability. In tropical Asia, investigation of the taxonomic status of the Iriomote cat is ofhigh priority, and an exception to this rule. The Iriomote
cat is the only felid taxon whose specific vs. subspecific(leopard cat: Category 5b) status is a matter of strong
debate(seeSpeciesAccount).
Identifying felid subspecies
and mappingtheir distribution will require a high degree of cooperation between
field biologists,systematists,geneticists,andmuseumand
zoo personnel. Genetics and morphology can be studied
from zoo animalsand museumspecimens. Specimens,
however, are scatteredamonga number of museums,andthereis no centralizeddatabaselisting felid specimensheldby the world’s various museums. Collection of samples
from captive animalsalsoposesproblems. First, asshownin Table 4, Chapter 5, over 60% of the officially catalogued captive populations of 20 cat speciesconsist ofgeneric animals,or animalsof unknown origin. Second,
captive-bred animalsare likely to be poor representativesof the genetic diversity found in wild populations. For
example, Miththapala (1992) notesthat it was difficult toanalyze the validity of the South China leopard on thebasisof samplestaken from captive animals:the popula-tion had beendeliberately inbred over severalgenerations
to maintain melanism.
Samplesfrom wild animalsare of critical importanceto
the task of mapping intraspecific diversity. This chapter
calls for a major increasein researcheffort to map species
meta-populations.In the processof carrying out surveys,
researchersshould coordinatewith systematistsand molecularbiologiststo ensurethat the biological samplesnecessaryto evaluate population uniquenessare collected.
Appendix 2 contains a field protocol for collection andstorageof suchsamples.

Genetics
Molecular researchcan addressfive important questions
relevant to conservation:(1) the uniquenessof speciesrelative to otherswithin the samegenusor higher taxonomic
unit; (2) the interrelatednessof speciessub-populations,leadingto understandingof historic patternsof movement
and the degree of gene flow; (3) the amount of genetic
diversity within a species,subspecies,or population; (4)the degreeof hybridization between individuals from thesameor closely relatedspecies;and (5) the breedingstructure of a population (Wayne et al. 1992). This field has
seengreat progressover the 1980sin both technique andapplication. Use of diagnosticgenetic fragmentsto determine degreeof relatednessbetween individuals, known
as DNA fingerprinting, was only recently developed(Jeffreys et al. 1985), andhasbeenrapidly improved andsimplified (reviewed by Wayne et al. 1992). One substantialimprovement wasthe developmentof polymerase
chain reaction(PCR), alsoin 1985,which permitsmultiple
copies of genetic material to be madefrom just a minute211Part II: Major

Issues.

Chapter

3. Research

Table 3
Genetic Research: Questions, Applicable Techniques,and Examples of Studies of Felids (modified from Wayne &a/. [1992])
QuestionsTechniques___-_______. -__________-._--__Allozymes
Karyology

1. Phylogenetic distinction

1

-P-P
mtDNA

VNTR

X

X

?

4-18

19-20

11-I 2,20-24

2. Phylogeographic history
3. Genetic variability
4. Hybridization
5. Breeding structure
Studies*

I-3

1 Explanation of techniques:
Karyology: Comparative analysis of the diploid number and morphology of chromosomes.
Allozymes: Comparative analysis of polymorphism at specific gene loci by protein electrophoresis.
mtDNA: Comparative analysis of divergence of gene sequences in rapidly evolving, maternally inherited
mitochondrial DNA.
VNTR: Comparative analysis of variable number of tandem repeat DNA (DNA fingerprinting).
2 Examples of genetic studies of felids:
1. Phylogenetic distinction: I. Wurster-Hill and Gary 1975; 2. Wurster-Hill and Centerwall 1982; 3, Wurster-Hill
ef al. 1987; 4. Collier and O’Brien 1985; 5. Wayne et al. 1989; 6. O’Brien this volume; 7. O’Brien ef al. 1987c;
21. Olmsted et al. 1992; 22. Janczewski ef al. 1992.
2. Phylogeographic history: 7,8. Yukhi and O’Brien 1990; 9, O’Brien et al. 1987a; IO. O’Brien et al. 1987b;
11. O’Brien ef al. 1990 and Roelke et al. 1993; 12. Packer et al. 1991 b; 19. Miththapala 1992; 20. MenottiRaymond and O’Brien 1993.
3. Genetic variation: 7-12, 13. Newman et al, 1985; 14. O’Brien et al. 1985; 15. Goebel and Whitmore 1987;
16. Miththapala et al. 1991; 17. Randi and Ragni 1991; 18. Hubbard et a/. 1992; 19-20,23. Gilbert et al. 1991.
4. Hybridization: 17-I 8.
5. Breeding structure: 12, 20, 23,24. Packer et al. 1991a.

its relevance to conservation biology.
The role of molecular research in understanding evolutionary relationships at the species level is discussed by S.
O’Brien under Taxonomy at the beginning of this book.
Hybridization between domestic cats and wildcats is discussed in the Species Accounts, and the question of subspecies was examined previously
in this chapter.
Molecular analysis has been used at a resolution finer than
the subspecies level by Packer et al. (199 1b) to examine
the relationship between two closely situated lion populations. Allozyme and DNA fingerprinting analysis was

initial sample. Even small bits of bone, skin, or hair frommuseum specimens can be used. These techniques, forexample, were used to analyze cellular DNA from theextinct saber-toothed cat Smilodon fatalis (of Ranch0 La
Brea tar pit fame), which the results suggest was a primitive relative of the modern pantherines within the mostrecent felid radiation (Janczewski et al. 1992).
Table 3 indicates the techniques that are appropriate to
the five areas of research outlined above, and reviews theirapplication to studies of felid genetics. Wayne et al.
(1992) provide a concise description of each technique and212

Par? II: Major

used to show that the small population of lions living
within Tanzania’s Ngorongoro Crater was probably
founded by immigrants from the adjacent Serengeti
National Park. The remainder of this discussion centers on
the importance of studies of genetic diversity and breeding
structure to felid conservation.
Genetic variation is thought to be essential to the longterm adaptability and persistence of populations by providing different options on which natural selection canoperate in response to environmental change. Although
the evolutionary significance of genetic diversity and theneed for its conservation is widely recognized andaccepted, actual demonstrations correlating genetic diversity and fitness have been few. Most studies have beenlaboratory experiments which examine only one index of
fitness (e.g., growth rate) (Beardmore 1983, Allendorf andLeary 1986).
Loss of genetic diversity within a population occurs
when it shrinks to a small size. This increases the chancesfor expression of harrnful genes. Expression of deleterious
mutations is synonymous with inbreeding depression,
which also refers to the consequences of matings between
closely related individuals. The physiological impairments resulting from inbreeding depression have been best
documented by breeders managing small captive populations, especially domestic animals (Ralls and Ballou
1983). However, while the potential of inbreeding to drive
isolated populations to extinction has been demonstrated intheoretical models (Senner 1980, Gilpin and Soule 1986),
in practice the costs of inbreeding, even in captive populations, have been difficult to predict (Ralls et al. 1988). In
other words, the exact consequences of loss of genetic
diversity are probably situation-specific,
and will vary
between both species and populations.
New evidence in support of the need to conserve large
populations in order to maintain genetic diversity may wellarise from future application of DNA fingerprinting to thebreeding structure of felid populations, as was done for
Serengeti and Ngorongoro Crater lions (Gilbert et al. 1991,
Packer et al. 1991a). Results showed that reproductive
success is highly skewed in coalitions of male lions: only
two males per coalition in the Serengeti fathered almost allof the offspring in their pride, regardless of the size of thecoalition (up to nine males). Not all potential breeders
have an equal chance of passing on their genes, and thus
lion populations are not panmictic (randomly mixed), as
is often assumed in simulations which model population
viability. Non-random breeding success leads to more
rapid decay of genetic variation than predicted by simulation models (Lacy 1993). So far this study has been the
only one to employ DNA fingerprinting to clarify thesocial organization and reproductive patterns of a population of wild felids, although captive populations have been
analyzed (e.g., Menotti-Raymond and O’Brien 1993).

Issues.

Chapter

3. Research

It appears to be a typical pattern of felid dispersal thatfemale offspring establish home ranges close to theirmother’s range. Based not on genetic analysis but on
known relationships, Smith et al. (1987b) found that the
average degree of relatedness between neighboring female
tigers in Nepal’s Chitwan NP was 0.35, similar to a typical value for a lion pride, which almost always consists ofclosely related females (Sunquist and Sunquist 1989). The
general pattern of felid social organization is that a male’s
home range overlaps those of several females and, if males
do indeed actually mate with all females within theirranges (molecular analysis is necessary to confirm this),the potential for inbreeding and more rapid loss of genetic
diversity arises. This is particularly the case if dominantmales prevent others from breeding and sire a disproportionate number of offspring, as has been found for tiger(Smith and McDougal 1991) and lion (Packer et al. 1991a)
populations. If the population is so isolated as to bar emigration of related and immigration of unrelated individuals, there is real cause for worry about population viability(see below). Smith et al. (1987a) suggest that this is a serious problem for tigers on the Indian subcontinent, where
the structure of the meta-population is characterized bypopulations that are either completely isolated, or have aprobability of genetic exchange considerably less than one
individual per generation.

Population

Viability

Analysis

A minimum viable population (MVP) is one that meets“the minimum conditions for the long-term persistence
and adaptation of a species or population in a given place”(Soule 1987a: 1). It is theoretically sufficiently large toprotect against extinctions caused by harmful and unpredictable genetic, demographic, or environmental factors
over a given period of time (generally expressed in hundreds of years). Determination of a generic, rule-of-thumb
MVP size has been the subject of considerable effort anddebate (reviewed by Shaffer 1990) and, based solely on
long-term conservation of genetic diversity, should beconsidered equivalent to an effective population size (Ne)of several hundred (Soule and Simberloff 1986, Lande andBarrowclough 1987).
The concept of effective population size summarizes
the genetic influences acting on a particular population,and has important implications for evolutionary processes(Wright 1969). Ne is defined as the size of an ideal population which maintains the same genetic diversity as thereal population (Kimura and Crow 1963), and is equivalent
to the number of breeding animals per generation. It is afunction of social organization and population demographics, and as a standardized measure permits comparison between species, between populations of the same213

Part II: Major

Issues.

Chapter

3. Research

Table 4
Ratios of Effective Population Size to Actual
Population Size (Ne/N) Calcu lated for Cat Populations
Speciesand Population

Ocelot, central Venezuela
Tiger, Chitwan National Park, Nepal’
Florida panther, U.S.*
Puma, southwestern Alberta, Canada3

Ne/N ratio

Reference

0.37
0.40
0.25-0.5
0.64

Ludlow and Sunquist (1987)
Smith and McDougal(1991)
Seal et al. 1989
Dueck (1990)

1 included good field data on a key component of effective population
reproduction, gathered over 17 years of field work.

size, variance in individual

lifetime* Upper estimate (0.5) based on proportion of adults known to have bred over 7 years of fieldwork. These
adults were treated as a single generation because not all animals were foilowed for entire period, andtherefore data is lacking on individual production of offspring. Lower estimate is speculative.
3 Dueck (1990) applied the formula developed by Reed et al. (1986) to field data collected by Pall et al. (1988).
In their assessment of estimators’ effective population size, Warris and Allendorf (1989) found that the formulaeof Reed et a/, (1986) tended to overestimate Ne/N by roughly 60%.
Part of the reason that there have not been more attempts to estimate Ne for cat populations is that reasonably
accurate demographic data are needed, of the type that can be collected only through long-term study, and thenwith difficulty. Harris and Allendorf (1989) suggest the use of population simulation models-basedon an initialshort-term series of field data as well as the willingness of biologists “to make educated, insightful guesses” for key
parameters related to social structure -to project population dynamics over individuals’ lifetimes. For management
purposes, they point out that it is probably not neccessary to strive for great precision in Ne estimates, given that
unpredictable variation in demographic or environmental events can easily alter the rate of genetic toss.

review in Chapter 1). However, MVP size decreases
sharply if the population is not completely isolated, butmaintainseven a low rate of genetic migration from otherpopulations. Simulation modelsindicate that immigration of new animalscan substantiallyreducea population’s
extinction risk (Soul6 1987c, Beier 1993). Many mammalshave beenshownto have low levels of genetic variation relative to other taxa (Selanderand Johnson 1973,
Powell 1974, Nevo 1978). Lande (1979) and ChepkoSadeet al. (1987) suggestthat low levels of genetic variation and chromosomalevolution patternsindicate that thegeneraltrend of mammalianpopulation organization is insmall,semi-isolatedandrelatively inbreddemes(sub-populations), occasionally augmentedby immigration fromneighboringdemes.For thesereasons,andbecausepumas
in fragmented habitat in the U.S. have actually moved
through narrow, natural landscapefeatureswithin developed areasto reach larger habitat patches(Maehr 1990,
Beier 1993), the needfor movement corridors to connect
smallisolatedreservescontaining big cat populationshas
beenemphasizedin Chapter 1.

species,and between the samepopulation at different
times. It is usually only a fraction of actual populationsize(N), becausenot all animalsin the population arebreeders.
The smaller the ratio of Ne to N, the greater the chancefor genetic drift and the greater the level of inbreeding
characterizing a species(Chepko-Sadeet al. 1987). Ne/N
ratios appearto have beencalculatedfor just three cat populations (Table 4). They imply that a minimum viable
population of several hundred breeding animals would
actually require nearly 1,000animals.
Moreover, a population size sufficient to mitigate environmental and catastrophic uncertainty (habitat change,or an epidemicor natural disaster)shouldbe considerablylarger than one required only to conserve genetic diversity (Shaffer 1987,Lande 1988). Taking thesefactors into
account, viable populationsare thus expected to be of theorder of several thousand individuals (Belovsky 1987,
Soul6 1987b,C. Thomas 1990).
It is clear that sucha goal is not obtainablefor populations of the bigger cats in protected areas. Most of theworld’s protected areasare simply not large enough(see214Part II: Major

However, considerable controversy surrounds the realsignificance and utility of the MVP concept, much of it in
the form of unpublished grumbling about whether such“ivory tower” thinking dictates that too-small populations
be essentially written off. There has been a great deal oftheoretical writing about conservation of MVPs, but few
attempts at application as a management practice (Shafer
1990, Caughley 1994). The correlation between genetic
diversity and fitness is theoretically robust but empiricallyweak (e.g., Beardman 1983, Allendorf and Leary 1986:
72-76), and would certainly benefit from further fieldwork. However, signs of inbreeding depression can bevery difficult to detect in the field unless the situation isvery advanced (Smith and McDougal 1991).
With regard to cats, there are three well-researched
examples which go furthest toward demonstrating small
population vulnerability in the wild, rather than just on
paper. The first, the Florida panther, is a very small population exhibiting relatively clear-cut symptoms ofinbreeding depression. The total population now numbers about 30-50 adults in fragmented habitat (Logan et al.
1993), and incestuous pairings have been observed(Roelke et al. 1993). Genetic diversity within the population is very low (O’Brien et al. 1990, Roelke et al. 1993).
Males have high levels of abnormal sperm and have overthe last 20 years suffered an increasingly and unusuallyhigh incidence of cryptorchidism(one or both testicles
undescended, with progressive loss of spermatogenesis).
A recent increase in cardiac abnormalities, in some cases
fatal, has also been observed (Roelke et al. 1993). An
analysis of the population’s viability concluded that theprobability of extinction within 20 years is high unlessthere is management intervention (Seal et al. 1989).
The second example illustrates the vulnerability ofsmall populations to unpredictable environmental events.
The population of lions in Tanzania’s Ngorongoro Crater
over 1957- 1961 was estimated to number 65-70 individuals, including young. In 1961- 1962, there was an infestation of biting fly (Stomoxys calcitvans) in the Crater,
following exceptionally heavy rains. Lions appeared tobe the preferred hosts for the flies; repeated bites led to skininfections and eventually emaciation. Unable to hunt
effectively, the population crashed to nine adult females
and one male by June 1962. Although the population
rebounded to its present level of between 75- 125 animals,
all members are descended from only 15 founders, andshow a significant lack of genetic diversity and high levels of abnormal sperm relative to lions from the nearby
Serengeti, the source of the founders. There are indicationsof declining reproductive success. The Crater is an ecological island which contains only a subset of the large
mammals found in the Serengeti. Wild dogs occurred in
the 1960s but have since disappeared. Male lions haveoccasionally immigrated into the Crater, but none have

Issues.

Chapter

3. Research

successfully bred since 1969 (Packer et al. 199 1b).
The third example shows why these populations should
not be written off. As discussed in the species account,wild cheetahs from both east and southern Africa are virtually identical in nuclear coding DNA, on a par withdeliberately inbred strains of laboratory mice (O’Brien etal. 1985). The cheetah population appears to have passed
through a severe population bottleneck around the time of
the late Pleistocene extinctions of large mammals(Menotti-Raymond and O’Brien 1993).
Yet the consequences for the cheetah today are notclear. Since the time of the bottleneck, there has been
ample time for natural selection to eliminate the most deleterious genes arising from inbreeding depression (i.e.,
those individuals with severe defects would not have survived). However, while moderate levels of variation haveaccumulated in non-coding genetic material, it would theoretically take “millions of years” to restore similar levelsof coding DNA heterozygosity for a species as depauperate in variation as the cheetah (Menotti-Raymond
andO’Brien 1993). Problems symptomatic of inbreeding
depression have been found in the global captive population, such as low levels of conception and elevated cub
mortality (Marker and O’Brien 1989), but the outstanding
reproductive successes achieved by some institutions indicate that inappropriate management is at least partly
responsible (Lindburg et al. 1993, Marker-Kraus
and
Grisham 1993, Wildt et al. 1993). Similar evidence ofpoor reproduction has not been found in the intensively
studied wild population in the Serengeti (Laurenson et al.
1992), despite Serengeti males having high levels ofabnormal sperm (Wildt et al. 1987a). Cub losses in the
Serengeti, which are very high (95%), are attributable toenvironmental factors, including lion aggression; moreover, females will rapidly conceive again following loss ofa litter. In reserves, cheetah numbers appear to be affected
far more by interspecific competition with other large
predators than by side-effects of genetic homozygosity(Caro and Laurenson 1994).
Because of their ecology, the big cats are highly vulnerable to isolation in small vulnerable populations. The
standard practice of setting aside a scattered network ofreserves may not prove to be very effective for these
species. The theoretical tools for analyzing population viability are available, as is the requisite field-collected biological data (on at least a crude level) for most big cats.
Better use should be made of tools such as GIS habitat
maps and population simulation models to demarcate subpopulations of these species, especially those of Categories
1-3, and a first-order evaluation of their status should be
done. Population viability analysis for cats has so far been
done for only three taxa- the Florida panther (Seal et al.
1989), the Sumatran tiger (Tilson 1992a), and the Asiatic
lion (Walker 1994). These efforts have been led by the215

Part II: Major

Issues.

Chapter

3. Researchin 1985 show that the population may have been exposed
to CDV around 1980. Since 1992, however, there havebeen four epizootics of CDV affecting predominantly
African and Asian cats in U.S. zoos. The extent of themorbidity and mortality in these recent outbreaks, coupled with the severity of the Serengeti outbreak, suggeststhat CDV has acquired increased pathenogicity for cats.
Controlling the Serengeti epidemic through vaccinationof the lions was not possible, as existing modified live
CDV vaccines were designed for domestic dogs and could
cause the disease when given to other carnivores.
The Tanzanian authorities launched a program in the
Serengeti area to vaccinate domestic dogs (the most likely
vector of the disease, since an outbreak of CDV occurredin the adjoining Masai Mara reserve in 1991). However,as shown by the 1980 outbreak of CDV in the Serengeti,
the disease can run its course, reducing but not threatening healthy populations overall, while vaccinating about
30,000 domestic dogs in the vicinity is a major challenge,
particularly since their average life span is 30 months andthe program would have to be continued for many years
to reduce the risk of new epidemics.
The rapid action of the Tanzanian authorities in carrying out an extensive veterinary investigation into the
Serengeti epidemic has provided important insights which
form the basis for continued monitoring of susceptible carnivore populations (L. Munson, C. Packer and M. Roelke,pers. comm.).
On the other hand, cats sometimes take advantage ofthe effects of disease upon prey populations. For example, wildcats in Scotland concentrate their predation on
wild rabbits afflicted with myxomatosis (Corbett 1979).
Although a wide variety of diseases has been documented for cats in captivity (e.g. Wallach and Boever
1983, Fowler 1986, Seidel and Wisser 1987, Pedersen
1988, Bush et al. 1992), opportunities for interspecifictransmission and host susceptibility are probably increasedin these conditions. For example, although Rasheed andGardner (198 1) isolated feline leukemia virus (FeLV)
from a captive leopard cat, it has not been reported fromany wild felid population except European wildcats, which
are closely related to and interbreed with domestic cats,
which are frequently infected with FeLV. However, it isstill not clear whether FeLV is transmitted from domestic
cats to wild cats, or rather the virus is sustained withinwildcat populations (McOrist et al. 1991).
Table 5 presents a preliminary list of viral and bacterial pathogens reported in wild cats. In some cases, thepathogen is detected by sera reactivity to an antigen in thelaboratory, rather than by clinical symptoms observed
either in the field or by examination or necropsy. In allcases, however, the etiology and effect of the viruses on
their host populations are unknown.
This lack of knowledge stems from practical difficulties

Conservation Breeding Specialist Group (CBSG), which
has done similar conservation analyses for a wide range of
non-felid species. The tools of conservation biology need
to be applied more systematically to wild cat populations,with more active involvement of field specialists.
Future research should not only go toward identifying
which cat populations are viable and which are not.
Population viability analysis does not necessarily identify
the specific threats facing a population, nor lead automatically to management solutions.
As emphasized by
Caughley (1994): if a population is declining or small, it
is more effective in the long-term to provide a cure (findways to increase population size) rather than merely treat
the symptoms (manipulate the population genetically).
The limiting factors, or agents of decline, should be identified through an experimentally-designed
study, andappropriate management actions taken. It will not always
be possible to increase population size-for
example, if
the limiting factor is the size of a reserve isolated in a hostile environment-but
it is important that research projects
are designed to address and, if possible, manipulate andalleviate the factors which led to the population vulnerability in the first place.

Infection

and Disease

There is increasing appreciation of the effects that infection and disease can have on wild populations (May 1988).
Infection refers to the presence of parasites (from bacteria
and viruses to arthropods) within a host; disease results
only if infection is clinically harmful. Infection and disease can potentially impact survival, reproduction, dispersal, and distribution of host populations (Scott 1988), as
well as their level of genetic diversity (O’Brien andEvermann 1988).
The actual effects of disease upon wild cat populations
are little-known. There is little documentation of the incidence of disease, let alone the impact, and a review of theliterature produced few examples of infection or diseaseleading to a major decline in a population of wild cats
(Table 5).
The most important and dramatic disease outbreakoccurred at the time of writing, so that it is not yet possible to draw conclusions about its impact. By July 1994, an
epidemic of canine distemper virus (CDV) had affected
20%-30% of the 3,000 lions in Tanzania’s Serengeti
National Park, and about 87 lions in a monitored population of 250 died or disappeared. The first notable clinicalsigns were facial and foreleg spasms, loss of control oflimbs, and seizures. More than half of all lions with clinical signs died or disappeared. Canine distemper has been
considered rare in felids, only occasionally affecting individual cats in zoos, but blood samples from Serengeti lions

216

Par? II: Major

Table 5*
Exposure to Infectious

Micro-pathogens

Documented

Issues.

Chapter

3. Researchin Wild Cat Populations

Pathogen

SpeCieS

AfSGl

Ref.

Anthrax@

CheetahNamiblac

15

Bubonic plague

BobcatNew Mexicu

8

Canine distempter virus (CDV)aj b

lion

Serengetic

20

Feline caliciviius (FCV)aPuma
VVjfdcat

FkXidaFrance1213Feline coronavirus (FCuV)aCheetahPuma
WildcatSerengeti
FtoridaFrancel&l7
1213Feline cytauxzoonosis

BobcatPuma

North America
Florida7
12Feline immunodefiency- type (FIV) viruses

lion
Cheetah
Puma
BobcatLeopardKruger, Serengeti
Serengeti
North America
FloridaKruger NP, So. Africa

12
I
I
I
19

Feline leukemia virus (FeLV)aWildcatWildcatScotlandc
France313Feline panteucopenia virus (FPV)aCanada lynx
BobcatPuma
Puma

North America
Caiifornia
North America
Florida49II12Feline rhinotracheitis (FHV)a

BobcatWildcatWildcatFloridaScotlandFrance12313Feline syncytia-forming virus

Puma

Florida12

RabiesaJ)

Canada lynx
BobcatEurasian lynx
Tiger
Puma
Puma
Leopard
Snow leopard

North America
Virginia
Slovakia *i
India
North America
FloridacCaucasus Mtn. region
Kazakhstan

4
5
16
6
14
12
18
18

a Pathogen potentially fatalb Generally considered uncommon in felids
CCat death ascribed to this pathogen
I. Ufmsted et al. 1992; 2. Spencer ellal. in press; 3. McOrist ef al. 1991; 4. McCord and Cardoza 1982;
5. Carey and McLean 1978; 6. Burton 1950; 7. Kier et al. 1982; 8. Poland et al. 1973; 9. Lembeck 1978;
IO. Heeney et al. 1990; I 1. Hansen 1992; 12. Roelke et a/. 1993; 13. Artois and Remond 1994; 14. Storer 1923;
15. Lindeque et al. in prep.; 16. Hell 1992; 17. Evermann et a/. 1993; 18. Heptner and Sludskii 1972;
19. Brown et al. 1993; 20. Anon. 1994e.

217

Part II: Major

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Chapter

3. Research

Lion stricken

by seizure

causedby caninedistempter

virus in the Serengeti,

Tanzania,

February

1993.

A proper disease monitoring program for small populations allows the best chance for successful intervention.

in the field, both in identifying sick animals and finding
dead ones, and then attributing with certainty the death todisease. Most wild animals are host to some form of parasite or another, and parasite loads can increase when the
animal is in poor condition due to other factors, resulting ina vector between infection and loss of fitness.
A program to monitor infection in wild populations ismost useful if carried out concurrently with a field study.
Thus, the movements, reproductive activity, and ultimate
fate of the study animals can be evaluated in the light ofknowledge of the presence or absence of infection. Whileclinico-pathological investigation remains the best way toidentify infection and monitor the role it plays within a particular population, it is also useful to collect blood samples
from study animals immobilized for marking or radio-collaring (see Appendix 2). Sera extracted from the samples
can be frozen and later tested for reactivity to various antigens, and can be used as well for molecular analysis. Skinand fecal samples can also be collected for examination for
macro-parasites.
M. Artois (in Zitt. 1993) suggests three reasons why
monitoring disease in wild populations is important anddeserves more emphasis from field biologists:

Even for larger populations, disease can be a major
environmental influence. Infectious disease can affectpopulations in a manner independent of host density
such that sparse and widely dispersed populations arenonetheless at risk.
Biological samples are important for building up clinical and ecological knowledge of the circulation ofpathogens.

Summary and Conclusions
This review of research effort for cats suggests that the following topics are of conservation priority. Areas for future
research are summarized generally below, with links to
specific priority projects in Part III.
1. Natural history studies of cats ranked in Categories l-3
are urgently needed (see priority projects in Part III).
Without basic information on behavior and ecology,
attempts to conserve these species are bound to be
somewhat inappropriate and inefficient. These studies
should be carried out both within protected areas, toprovide baseline data, and in modified habitat typical of

1. Disease can have a devastating impact on small populations, as described for the lions of the Ngorongoro
Crater (Packer 1991b), or as in the well-known case of
the black-footed ferret (Thorne and Williams 1988).

218

Part II: Major

Issues.

Chapter

3. Researchand 103). They are most suited to protected areas
where the population dynamics of other non-felid
species, especially prey, are also studied. In the future,long-term studies should be mounted in protected areas
which hold large populations of cats of conservation
concern, on either a regional or global level. It is also
important to study the predator-prey dynamics of cat
populations located outside protected areas. Such studies could most appropriately be undertaken by having
a field researcher work under a long-term development
project with a conservation component.

the predominant regional form of land use, to assess
implications for species ‘conservation.
2. Most of these species occur in tropical Asia and Latin
America. In addition, the cats of north Africa andsouthwest Asia have scarcely been studied. Natural
history studies should be carried out with the active participation of local residents and experts, including
hunters. The value of these studies would be increased
if several cat species were studied simultaneously by asmall team.

6. Research projects designed both to develop and implement solutions to conflict between big cats and people
are of high priority (see Chapter 2, and Projects 5,6,22,
23,3 1,32,5 1,52,71,74,93,
and 102). Studies should
be carried out in areas typical of the predominantregional form of land use, or should alternatively focus
on potentially viable, unprotected populations.

3. Small, isolated populations are highly vulnerable toextinction. This has been well documented for cats,
particularly the large species, yet there has been little
systematic effort to map out cat population fragmentation and distribution beyond the more obvious examples of occurrence in protected areas. A number ofpriority projects have been put forward in Part III to
conduct presence/absence surveys to map discrete populations of cat species. Particular attention should be
paid to identification of potential and actual habitat corridors to permit migration between otherwise isolated
populations.

7. There have been relatively few attempts to specifically
apply the lessons of theoretical conservation biology
to the field. A cooperative effort should be made to
define Minimum Viable Population size for cat species,
based specifically on insight gained through long-term
studies of behavior and ecology (see Project 8). This
exercise would greatly aid reviews of species status,and prioritization of conservation effort. The following
species should be looked at first: Iberian lynx, tiger,
snow leopard, cheetah, lion, jaguar, puma, leopard.

4. Standardized, replicable techniques to census cat populations still need to be worked out, especially for thetiger in India (Project 48). Otherwise, there is little
hope of accurately assessing the effectiveness of population conservation efforts. Density estimation methods(spoor surveys, photo traps) should be tested against
populations whose size is known from long-term study(Project 7).

8. Field studies of cat populations could benefit fromcloser cooperation with the laboratory sciences. Field
researchers should take biological samples from cat
species at every opportunity to help advance knowledge of felid taxonomy, genetics, and disease (seeProject 9).

5. Long-term studies are necessary to understand “typical” felid population dynamics and the role played bycats in ecosystems (see Projects 16, 17,24,30,45, 101,219

Part II: Major

Issues.

Chapter

4. TradePart II
Major Issues in Cat Conservation

Chapter 4
Trade
For example, in the 1989- 1990 season the average pricepaid to the trapper for a Canada lynx pelt was CA $117
(U.S. $85), which is high compared to the average value
for other fur-bearers-U.S.
$67 for fisher, $57 for marten,$17 for bea ver, and $9 for fox. In 1984-1985, lynx pelts
were much more valuable-at approximately U.S. $445,
worth 18 times as much as a fox pelt (Canadian Wildlife
Service in Zitt. 1994). In other words, relatively low harvests can generate significant revenue. Theoretically, furtrade has the potential to be an important component of asustainable use strategy for cat conservation, providing an
economic incentive to maintain wild lands. The management of commercial trapping of the two North American
lynxes is reviewed to examine whether such an approach
actually works.
Illegal trade, on the other hand, has the potential to beamong the fastest and most destructive threats to wild catswhen it leads to commercial poaching (as opposed to illegal hunting for other reasons, such as livestock protection).
The fur trade from Latin America in the late 1970s to early
1980s is a good example. The region was the major supplier to the world market, but most exports were in contravention of national hunting and export bans. Imports up
to the mid-l 980s were legal, however, so that import statistics indicate how large the illegal trade was. When trade
is clandestine and “underground,” however, it can be very
difficult to gauge its true volume, identify consumer motivation and appropriate corrective measures, and enforce
prohibitions effectively. When the product is high-value,as most cat products are, the resulting environment is conducive to official corruption, which further compounds
efforts to bring the problem under control. The dynamic
depends very much on the product and its market. Illegal
trade in cats and their products is reviewed, and the conclusion reached that it is at its worst with regard to the big
Asian cats, particularly the tiger.
This chapter reviews the history of the international
trade in cat furs and other products up to the present time.
Trade in cat products cannot be evaluated separately fromhunting, and the biological impact of commercial hunting

Introduction
Historically, cats have been most prized for their fur. The
glamour of cat fur can probably be traced back to the firstpeople brave enough to slay a fearsome large predator. In
many cultures around the world, cat pelts are still worn by
the elite as symbolic expressions of status and power.
These overtones, coupled with the physical beauty of cat
fur, also explain their appeal to the fashion-conscious modern woman. An advertisement typical of the 1960s and1970s enticed buyers with the following: “Untamed.. .theSnow Leopard, provocatively dangerous. A mankiller.
Born free in the wild whiteness of the high Himalayas only
to be snared as part of the captivating new fur collection...styled and shaped in a one-of-a-kindness to bring out
the animal instinct in you” (Conway 1968).
The appearance of large numbers of pelts of tigers, leopards, jaguars, snow leopards, and their kin in boutiquesfrom Montevideo to New York and Berlin led to alarm that
trade might drive these species to extinction. This was
one of the primary concerns which fueled the developmentof the Convention on International Trade in Endangered
Species of Wild Fauna and Flora (CITES-see
Box l),
which entered into force in 1975. As implementation of
the Convention by its signatory states (particularly theimporters) has improved, those species of spotted cat
traded in large numbers, both before CITES entered into
force and in its early days, have now almost completely
disappeared from the fur trade. The volume of cat skins
legally traded in 1990 is only about a quarter of the volume of trade in 1980 (WCMC unpubl. data). Latin
America and sub-Saharan Africa have been replaced as themain suppliers by four northern countries-theUnitedStates, Canada, China, and Russia. The modem fur trade incats relies almost entirely on three lynxes and the Chinese
leopard cat.
Cats are one of the wildlife families used by the furtrade which are entirely wild-caught, rather than farmed.
Their pelts are also high-value compared to most otherspecies in the trade, even when demand is relatively low.

220

Part II: Major

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Chapter

4. Trade

Box 1
The Convention
Conve ntion on International Tra
Tradede in Endangered
Species of Wild Fauna and Flora (Cl(CITES)
TES)
lbred specimens, which can be traded commerciaHy).
Both import and export certificates are required before aninternational transaction can take place* Trade in
Appendix li species is supposed to be permitted only
when the exporting country’s Scientific Authority (typically the government agency responsible for wildlife
management) has determined trade to be “not detrimental to the survival of the species”‘. Trade in both Appendix
II and ill species require export permits only. The three
Appendices are amended at bi-annual conventions bymajority vote of the Parties, ostensibly in response tochanges in the conservation status of a species or subspecific populations (although sometimes politics overwhelms biological considerations). AlI fetid species arelisted on either Appendices l or II; the listing as of March
1995 appears below. New listing criteria were approvedat the Conference of the Parties in November 1994
because the previous criteria were not considered sufficiently objective or biologically based. Some cat species
listed on Appendix I cannot seriously be considered to beendangered by trade.

CITES is a treaty which restricts and regulates trade inwildlife. Its preamble recognizes that “international cooperation is essential for the protection of certain species ofwild fauna and flora against overexploitation throughinternational trade.” It entered into force on July 1,1975,
with a total of IO nations as ratified signatories. As of
May 1994, its Parties (signatory nations) number 122.
CITES functions on the basis of its four appendices,establishing different trade restrictions for species listedon Appendices 1, 11,and IIl and allowing for trade only
when a competent government authority has issued apermit conforming to the permit form in Appendix IV(8Gutigam 1989). Species are listed on Appendices i or
11by majority vote of the Parties.
Appendix I: includes “‘all species threatened with extinction which are or may be affected by trade. Trade in
specimens of these species must be subject to particularly strict regulation in order not to endanger further theirsurvival and must only be authorized in exceptional circumstances.”

In theory, the treaty permits an unprecedented degree ofcontrol over international commercial trade in wildlife,and should have far-reaching effects toward assuring
the viability of threatened species populations aroundthe world. Moreover, a comprehensive global tradeoversight mechanism is established through a requirement that signatories submit annual reports detailing alltransactions which require CITES permits (import, export,and re-export). Data from these reports are managed
by the World Conservation Monitoring Centre (WCMC)
in the U.K. under contract to the CITES Secretariat in
Switzerland. In practice, however, a number of difficulties impede both the functioning of the treaty and thecompilation of accurate trade data; the following of which
are most pertinent to the commercial trade in cats andtheir products (Johnson 1989).

Appendix If: includes “(a) alf species which atthough notnecessarily now threatened with extinction may become
so unless trade in specimens of such species is subject
to strict regulation in order to avoid utitization incompatible with their survival; and (b) other species which mustbe subject to regulation in order that trade in specimensof certain species referred to in the above sub-paragraphmay be brought under effective control” (i.e., so-called“look-alike” species which are difficult to distinguish fromothers listed on Appendix 1. Several felid species havebeen fisted on Appendix II for this reason).
Appendix 1H: includes “all species which any Party identifies as being subject to regulation within its jurisdiction
for the purpose of preventing or restricting exploitation,
and as needing the cooperation of other parties in thecontrol of trade.” Any Party may unilaterally list a native
species on Appendix If I.

1. Not all countries are Party to CITES. Trade between
these countries is not reported to CITES, and someostensibly illegal shipments may be “laundered”
through these countries.

in general, trade in Appendix I species is prohibited
except under certain essentially non-commercial circumstances (zoological exchange, scientific study, movement of personal effects or legitimately taken hunting
trophies, etc. -with the exception of certified captive-

2. The effectiveness of the implementation of CYTES
varies from country to country. Some have notContinued on next page.
227

Part II: Major

Issues.

Chapter

4. Trade

up in the annual reports of importing countries were nut
fisted in those of exporting countries, and vice versa.
For example, in 1978 West Germany reported theimport of 10,171 0nciJJa pelts from Uruguay but,aJthough Uruguay was the only Latin American country
tu submit a report for that year, the shipment of oncillapeJts was not recorded (Soydell 1980). Meanwhiie, inWest Germany’s 1979 CJTES report, no impurts wereJisted from Brazil, which had banned aJJexport of wildlife
and wildlife products, but officiaJ German Custums statistics reported the import of 61,4UU cat skins from thatcountry (Broad 1987). After 1980, correlation between
Germany’s import figures and others’ export figures
improved-andshowed that skins continued to beimported in Jarge numbers from Paraguay, which had
ostensibly banned the export of spotted cats in 1981
(CaJdweJJ 1984, Broad 1987). The effectiveness of theConvention depends heavily upon the Parties’ observance of national JegisJation, which can be strJcter thanthe CJTES Appendices.
WWF-US. is publishing aseries of guides to national wiJdJife trade legislation(Fuller et al. 1987, NichoJs et aI* 1991) which will servea very useful purpose.

enacted domestic legislation to impJement or conformto the treaty, in part or in entirety. Even one of themust scrupulous examples of implementing Jegislation, the Endangered Species Act of the UnitedStates, has its share of loopholes and red tape
(Kosloff and TrexJer t987, O’Brien and Mayr 1991).
Problems with forged CJTES documents occur frequentiy. The accuracy, frequency, and format of“annuaJ” reports (some Parties have never submitted
even one) are highty variabJe.
3. The scope of CITES is necessarily limited in that it
deaJs only with international, not domestic, trade inlisted species. The Convention has no direct impact
on a country’s domestic wiJdJJfeconservation policies.
4. Different Customs agencies use different units ofmeasure (e.g., kg of skins vs. numbers of pelts or skinplates), which hinders cJarity when WCMC compiles
reported trade data. Customs agencies may treat
country of origin differentJy (e.g., skins exported from
Paraguay through France to a third destination countG might be reported as originating in either Paraguay
or France, depending on the non-standardized criteria used by the destination country).

Despite its weaknesses, CJTES is an indispensable tooJfor controlling internationaJ trade in wiJdJife. While it onJy
works as weJJ as individual Parties, at the very least it
provides the best information available on gJobal tradepatterns for aJJspecies of cats and helps to fix an orderof magnitude on market sizes and organization.

5. Jliegal trade is obviously not reported through CITES.
During the early years of the Convention, these problems were acute. Shipments of cat furs which showed

Cat Species Listings on the CITES Appendices,

Larch 1994

Appendix I

Appendix If

Cheetah, A. j&at&
Caracal, C. caracaP
Asiatic golden cat, C. temmincki
BJack-footed cat, F. nigripes
Jaguarundi, H. yaguarondi3
Ocelot, 1. pardaiis
OnciJJa, L. tigri#us
Margay, L. wiedi
Jberian lynx, L. pardinus
Geoffroy’s cat, 0. geoffroyi
Andean mountain cat, 0. jacubitus
Leopard cat, P. 6. bengalen&
Flat-headed cat, P. planiceps
Rusty-spotted cat, P. rubiginosus5

Caracal, C. caracal
J3ornean bay cat, C. badia
Chinese mountain cat, F, bieri
Jungle cat, F. chaus
Sand cat, F. margarita
Wildcat, F. silvestris
Jaguarundi, H. yaguarondi
Serval, 1. serval
Canada lynx, L. canadensis
Eurasian lynx, L. lynx
Bobcat, L. rufus
Pampas cat, 0, colocolo
Kodkod, 0. guigna
Manul, 0. manui
Conthmd cm next page222

Par? II: Major

Appendix I

Issues.

Chapter

4. TradeI

NoteS

1 Annual export quotas fur live specimens and hunting trophies are
granted as follows: Botswana-5; Namibia-l 50; Zimbabwe-W.
2 Asian populations only.
3 Central and North American populations only.
4 Bangladesh, India, and Thailand pupulatkxw.
5 Indian population only.
6 Three subspecies: cut-@(Florida), costarieensis (Central America)
and cougar (eastern North America).
7 Annual expoti quotas fur hunting trophies are granted as fu~luws:IWswarta-100;
Central African Republic-40; Ethiopia-XXI; Kenya-W; Malawi-SO; Mozambique-6U;
Namibia-l 00; South Africa-75; Tanzania-25U; Zambia-300; Zimbabwe-500.

on wild cat populations is discussed. The relationship of
hunting and trade to cat conservation is examined bymeans of two case studies: the bobcat and lynx harvest
programs in North America, and the illegal Asian trade intiger bone.

International

The mushrooming popularity of spotted fur in North
America is traced to an appearance made in 1962 by theAmerican First Lady Jacqueline Kennedy wearing a leopard coat (Fig. 1), setting off a major trend (Nilsson et a2.
1980). The IUCN issued a warning at its 1963 General
Assembly that “the present fashion...of spotted cats is athreat to the continued existence of these kinds of animals,”with particular reference to the leopard and the jaguar(Anon. 1971). Estimates of annual imports into the UnitedStates and Europe during the late 1960s indicate that thepelts of over 10,000 leopards, 15,000 jaguars, 3,000-5,000
cheetahs and 200,000 “ocelots” (in this case, a trade namecovering similar species such as the margay and oncilla)reached the market annually (Gieteling 1972, Fitzgerald1989). A top quality jaguar coat fetched U.S. $20,000 in
the fur boutiques of New York, and Myers (1973) estimated that international trade in spotted cat skins wasworth approximately U.S. $30 million at this time.
In September 197 1, the International Fur Trade
Federation (IFTF), in association with IUCN and WWF,
concerned that the current level of trade might pose a threat
to some cat populations, strongly recommend to its members a voluntary moratorium on trade in the skins of threelarge cats-tiger, snow leopard, and clouded leopard-anda three-year ban on the leopard and the cheetah. The IFTF
also helped to fund investigations into the status of
exploited cats in South America and Africa; both studies
concluded that international fur trade had led to unsus-

Trade in Cat Furs

Records from the early days of the fur trade are scarce,
but for one year, 1763, there are records showing that
4,150 lynx pelts were exported from Canada to England,comprising a mere 2% of the furs in trade that year (Poland1892). However, records of purchases of Canada lynx
pelts by the Hudson’s Bay Company during the 19th century attest to the growing popularity of cat pelts. Peakharvests were of the order of 80,000 pelts annually in thelate 1800s but they declined sharply after the turn of the
century (Elton and Nicholson 1942). In the early 1900s
approximately 64,000 bobcat and lynx pelts were soldannually in the United States, the world’s largest fur market for much of this century (Osbom and Anthony 1922).
Following the Depression and World War II, the furtrade’s source of supply underwent a major shift frommainly wild-trapped to mainly ranched animals (IFTF
1989). Cats, however, are not ranched, and their proportion within the wild-caught
minority of furbearers
increased dramatically during the 1960s.

223

Part II: Major

Issues.

Chapter

4. Trade:igure1. Cat fur fashions

in the 1960s

tainable hunting pressure in many areas (Koford 1976,
Myers 1975, 1976). The IFTF’s moratorium and ban were
not universally respected (see Appendix 6 for a statementby the IFTF). Producer and consumer countries began topass protective legislation for many of the larger cats and,
when CITES entered into force in July 1975, all felids
were listed on either Appendix I or II (see Box 1). With all
the exploited big cats thus prohibited from internationalcommerce via their placement on Appendix I, the industry focused on the smaller species.
The small spotted Latin American cats had been tradedthroughout the 1960s as well, and in greater volume thanthe larger cats, in part because it takes more of them todisplayedin Animals

magazine.

Source:Anon.

(1971).

make a coat. “Ocelot” pelts comprised the vast majority,with 133,069 officially imported into the United Statesalone in 1969 (Myers 1973). As ocelot populations were
hunted out and publicity increasingly focused on the needto protect this species, trade in pelts of other small spottedcats of South America-themargay, oncilla, andGeoffroy’s cat-grew during the late 1970s to early 1980s.
Trade volume in felid pelts was very high during thisperiod, and was the subject of a special review in the 1978
annual report of the CITES Secretariat (Anon. 1978). The
report notes: “The available customs figures for all countries in 1977 total approximately 570,000 imports and
6 15,000 exports. Although there is a good deal of over-224

Part II: Major

lap in these figures, the true totals are much higher becauseof the lack of Customs’ data from most countries.”
Problems with documenting the extent of the trade in cat
furs lent impetus to calls for governments to be regular insubmission of annual trade reports, and to the creation ofa centralized data base to monitor that trade.
Many South American countries had prohibited trade in
their native cats before CITES became effective (e.g.,
Brazil in 1967, Venezuela in 1970, Chile in 1972,
Colombia in 1973, Peru in 1975, Argentina in 1976: Broad
1987). However, traders from these countries smuggledskins out and laundered their exports to markets abroad
through other outlets. Colombia’s Amazonian free portof Leticia was notorious (Smith 1976). Leticia’s status
was revoked in 1973 after the documentation of widespread abuse of trade privileges (Inskipp and Wells 1979).
It is apparent from the numbers exported that the bulk offurs exported from Paraguay and Bolivia during the period
1979-1985 must have been re-exports of pelts taken andsmuggled in from neighboring countries, which were formerly major exporters with trade networks in place. In
Europe, the free port of Hamburg facilitated the entry of
shipments with CITES documents either of dubious validity or lacking altogether, as management authorities had no
jurisdiction in these zones (Inskipp and Wells 1979).
Customs in some countries also had no jurisdiction overtransit trade.

1979

1980

Latin American

Figure 2. Trade
Reports.

in skins

1981

speciesof Felidae,

1982

1983

1984

1985

Lynx/Bobcat/Bengal
LeopardJ/(//////1
19784988.

Source:TRAFFIC

Chapter

4. Trade

Since then, the world trade in cat skins has shown asteadily declining trend, and is now largely legal. In 1980,
CITES-reported
world trade in cat pelts totalled some450,000; by 1990, it had fallen to just over 100,000(WCMC unpubl. data). The number of species in trade has
declined by more than half, and trade in the remaining
species is decreasing. A major cause of the decline was astrict import ban enacted by the European Community in
October 1986 on Latin American spotted cat furs, and on
leopard cat in 1988. For example, 82,500 cat skins legally
exported from Argentina in early 1987 were refused entry
into Europe (Broad 1987). Changes in fashion have been
another major factor, brought about in no small part by
the campaigns of pressure groups.
As exports from Latin America have fallen steeply inrecent times, legal trade in cat furs now involves only
countries of the northern hemisphere, and has depended
on just four species: the bobcat, Canada and Eurasianlynxes, and the leopard cat from China (Fig. 2). As withthe small neotropical spotted cats, these four species havebeen harvested in large numbers, and there is cause forconcern that trapping pressure in the 1980s on at least two
species, the Canada lynx and the leopard cat, has led topopulation declines (see Species Accounts for details).
Table 1 provides an overview of species in the skintrade. Numbers of pelts in trade (net trade) have been calculated from data in annual reports to CITES (McMahan

500

1978

Issues.

1986

1987

Cat

Total
- -w

international

from

225

CITES

1988

AnnualPart II: Major

Issues.

ChapterTable ICITES-reported

4. Trade.
International

High volume commercial

Trade in Cat Skins, 1976-l 990

trade, current (total 108,ooo - 1.3 mitlion skins)

Species

AverageAnnualTrade

Trade
in 1990

Major
Expurters~

Trendleopard cat, P. bengalens&
Bobcat, L, rufus
Canada lynx, L. canadensis
Eurasian lynx, L. /ynx

85,98557,20819,034
7,227

86,911
18,405
7,386
3,366

CN
US, CA
CA, US
CN, RU

Reclining
Declining
CyclicalHigh volume commercial

trade, past (total 76,000 - 470,UW skins)

Species

AverageAnnualTrade

Peak Years

Geoffroy’s cat, 0. geclffroyi
Oncilla, L. tig&ws
OceIut, 1. pardalis
Margay, L. wiedi
Wildcat, F. silvestris

31,107
23,584
14,548
8,548
5,125

1978-I 984; 1987
1978-I 984
1976-l 983
1977-l 983
1977-l 986

Low volume commercial
SpeciesJungle cat, F. chaus
Pampas cat, 0. colocolo
Manul, 0. manul
Caracal, C. caracal
Serval, L. servalPrimarily non-commercial,
SpeciesLion, P. lea
Leopard, P. pardus
Puma, P. concolor
Jaguar, P. onca
Cheetah, A. jubatus
Tiger, P. tigris

Trade
in 1990
1,521
0
21
2
46

trade, past (total 6,000 - 45,000 skins)
AverageAnnual
Trade
2,909
1,955
1,721
684
392

Peak Years

Trade
in 1990

1979-I 983; 1988
1979-l 981; 1987
1978-I 984; 1987
1980
Steady decline

0
0
0
35
6

especial ty trophies (total 200 - 10,000 skins)
AverageAnnualTrade

Trade
in 1990

645
566
294
117
102
16

688
657
70
0
8
9

%A = Canada; CN = China; RU = Russia; US = United States.

226

Trend

Slightly increasing
Slightty increasing
Steady, tow
Low, declining
Declining
Steady, lowPart II: Major

1986, Broad 1987, WCMC unpubl. data). Fifteen yearsof trade data have been averaged for each species to index
the representation of each within the skin trade.
The bobcat and leopard cat have been part of the furtrade in greater numbers and for a longer period of timethan the Latin American small spotted cats, even though
the latter have more good quality habitat available to them(Amazon Basin). However, concern among the Parties to
CITES regarding the biological impact of the fur trade has
centered on the neotropical cats, to the extent that four
species -the ocelot, oncilla, margay, and Geoffroy’s catwere upgraded from CITES Appendix II to Appendix I in
1989 and 1992, despite the fact that international trade inpelts had virtually ceased by the mid- 1980s.
In 1985, the Parties actually downgraded the Chinese
population of the leopard cat subspecies P. bengalensis
bengalensis, which is believed to occur only in Yunnan
province (J. Yu in Mt. 1991) from Appendix I toAppendix II, even though no management programs werein place and no status surveys had been undertaken (tworequirements upon which the Parties are usually insistent).
Exports of leopard cat skins from China shot up shortly
thereafter, from about 19,000 pelts in 1983 to 89,000 in1984 and over 200,000 in 1987. Based on a survey of theinventory held by major fur trading companies in China,
J. Yu (in lift. 199 1) believed that this subspecies made up
the majority of Chinese exports, having a tendency towarda richer, bolder pattern which is more commercially valuable. Increasing concern about possible overexploitation
led the CITES secretariat to call on member countries in
April 1993 to refrain from further imports of leopard catskins from China pending implementation of suitable control and management measures in that country. A projectis currently underway to evaluate the species status andcurrent management measures in China, and to advise thegovernment on necessary improvements (see Project 13
in Part III).

Issues.

Chapter

4. Tradeaffect cat populations. Canada lynx are particularly vulnerable to over-harvest when their main prey species,
snowshoe hare, go through cyclic declines. Thus, indicesof abundance are of critical importance in developing sustainable harvest regimes, but these have proved difficult
to develop for cats (see Chapter 3, Research).
Unfortunately,
the impact of the past 30 years ofinternational commercial trade will never be fully known
for most species. For the spotted cats of Africa and Latin
America, hunting was either illegal or unmanaged, so thatthe biological impact was never addressed. That none of
the species involved have become extinct does not necessarily mean that the trade was sustainable. Important subpopulations may have been eradicated-or
may not
have-but at this late stage this issue will be difficult, if not
impossible, to assess. Identification of areas subject toprolonged offtake remains very much local knowledge.
Numerous hunter and trader accounts refer to local depletions (Myers 1975,1976, Koford 1976, Smith 1976,
Hamilton 198 1, Melquist 1984, Martin and de Meulanaer
1988). Evidence of local recovery (such as the ocelot in
Venezuela: R. Hoogesteijn, pers. comm.) is anecdotal, andthe current situation in areas previously subjected tointense hunting pressure has not been investigated.
Moreover, the impact of habitat loss cannot be consideredseparately; populations that have been fragmented and isolated by human land-use patterns are particularly vulnerable to elimination.
Martin and de Meulanaer’s (1988) attempt to simulate
the impact of the fur trade on regional leopard populationsin Africa was a thought-provoking exercise. Using a variety of data sources, they estimated the number of leopards
killed annually in sub-Saharan Africa for the fur trade
between 1950 and 1986 and, according to a leopard population model they developed, examined how large leopard
populations would have had to be in order to survive theofftake. Figure 3 illustrates a first attempt to answer thequestion about the impact of this period of uncontrolledcommercial exploitation of cats. In the case of northern
Africa, the harvest was apparently so large that Martinand de Meulanaer (1988) had to double their estimate of
the potential leopard population existing at that time inorder to prevent its simulated extinction. However, theoverall conclusion was that, based on population estimates
predicted by their model, the fur trade had no serious lasting impact on the abundance of the species.
Only for two species-the
Canada lynx and the bobcat-has the biological impact of commercial harvesting
been a priority for study. These are relatively important
species in the fur industry, and programs to manage commercial trapping are in place. An examination of theeffectiveness of these management regimes is useful forassessing whether commerce can play a positive role in cat
conservation.

The Biological Impact ofTrade on Wild Populations
In general, it appears that cat populations are resilient toharvest up to a certain threshold (“maximum sustainable
yield”), and offtake in excess of that threshold can lead to
extirpation. The percentage of a population that can be
harvested without producing an overall decline in numbers
varies among species-for example, the bobcat has greater
fecundity than the ocelot and can sustain higher offtake.
Within a species, however, local environmental differences will lead to a variation in density across its range,
so that a harvest rate appropriate for one area may be inappropriate for another. Changes in prey abundance will

227

Part II: Major

issues.

OENS

kn)

I TY

(/me+

Chapter

4. Trade

*

OENSI

TY

(1-q.

km)

HARVEST

I. 01

1950

1960

1970

1980

1990

2000

1970

HARVEST:

100

AREAa

5 959 000 l q. km

POPULATIONI

145

SOUTHERN-

13 500000

lropordleopardAFRICA

1980
YEARS

YEARS

(LCI

-

mrtimoto)

RESPONSETO HARVESTHARVESTt

100

AREAt

3 544

000 l q. km

POPULATIONI

100

loopordNORTHERN-

14 SO0000

AFRICA-loopord(roviood

REVISEDl rtimoto)RESPONSETO HARVEST

Figure 3. Predicted
response
to harvestof leopard
populations
in southernand northern
Africa.
Source: Martin and de
Lesotho, Malawi, Mozambique,
Namibia, South Africa, Swaziland,Zambia,Meulenaer(1988).
[Southern
Africa = Angola, Botswana,and Zimbabwe;
Northern
Africa = Djibouti, Ethiopia, Somalia, and Sudan].

both the local and national levels, to ensure their conservation. Canada and the United States have developed themost advanced management programs to date for commercial exploitation of feline furbearers. The following
sections review North American management of comrnercial hunting of bobcat and Canada lynx, and examine towhat degree these programs can be considered (1) examples of consumptive sustainable use of cats and (2) to support the conservation of these species. Any government
which considers commercial exploitation programs fortheir cat species would be well advised to review thestrong points and weak points of North American programs in the process of developing locally appropriate
management practices.

Sustainable Use of Catsfor the Fur Trade: The
North American Example
It would be inappropriate to review large-scale commercial
trade apart from the context of harvest management policies (or lack of such) which make it possible. Basic components of the North American strategies are reviewed
below. Similar information on Russian management of
their Eurasian lynx harvest is not available, and China’s
high leopard cat harvest, with estimates of annual takebetween 1985 1988 as high as 400,000 (J. Yu in litt. 199 I),
is at present unregulated, although a project is underway toinvestigate biological impact and develop appropriate
management controls (Johnson and Fuller 1992, Johnson
et al. 1993).
Although the fur trade has posed a threat to some cat
species, in the case of others it can be argued that it benefitted their conservation.
The harvest and sale of felid
pelts can be a component of a strategy to extract economic
value from wildlands, providing a strong incentive, on

Review of Bobcat and Canada Lynx
Management
Programs in the UnitedStates and Canada
In the United States and Canada, bobcat and Canada lynx
populations are managed at the state, provincial and Indian228

Pat? II: Major

2. All pelts should be registered and marked;
3. Harvest level objectives should be determined annually.
In Canada, each province sets its own information andmanagement requirements. Quinn and Parker (1987) havesuggested that the minimum information requirements
should include the following:
1. The chronology of lynx harvest statistics and prevailingregulations relative to specific management regions;
2 The chronology of snowshoe hare abundance relative to
specific management regions (i.e., small game license
returns, questionnaires, etc.);

Information on total harvest of the species;
Information on geographic distribution of the harvest;

3. Assessment of the above to evaluate the magnitude andtemporal trends of the lynx-hare cycle at regional levels;

Habitat evaluation.

lynx, Yukon,

Canada,where

studies

4. Trade

1. Harvest should be controlled, the methods and seasons
to be a matter of state choice;

Population trend information, the method of determination to be a matter of state choice;

CanadaChapter

The minimum national requirements for bobcat management programs are:nation levels, rather than by the national governments. In
the U.S., the role of the national government is chiefly toensure that the total amount of furs exported does not constitute an offtake which is detrimental to these species’ survival, as required by CITES for trade in species listed inAppendix II (see Box 1). The Canadian government, on
the other hand, has delegated this responsibility to theprovinces (C. Dauphine in Zitt. 1994).
In the U.S., in the late 1970s the Endangered Species
Scientific Authority (the responsibilities of which werelater transferred to the Fish and Wildlife Service), established the following biological information requirements
to evaluate the adequacy of state bobcat management programs for export approval:

Radio-collared

Issues.

of fur-bearing229

animals

are being carriedout.

Part II: Major

Issues.

Chapter

4. TradeAnnual samples of lynx carcasses (collected from trappers) to assess sex and age’structure and fecundity;

and educated guesses (Bluett and Tewes 1988).
While varying degrees of effort go into evaluating cat
habitat, it is rarely actively managed for the cats, despite anumber of specific recommendations having been put forward (e.g., Allen 1987, Quinn and Parker 1987, Koehler
and B&tell 1990). Generally speaking, cat habitat is managed in a variety of ways by a number of state, provincial,and national agencies with diverse briefs (i.e., state
wildlife management areas, national forests, nationalparks, watershed conservation areas). Even where habitat
management guidelines have been developed and officially approved, as for Canada lynx in Washington state(Anon. 199 le), implementation is spotty due to the factthat not all agencies whose jurisdictions include cat habitat
are bound by them (S. Thorpe in Zitt. to USFWS 199 1).
An example of recommendations for specifically managing habitat for cat conservation is Koehler and B&tell(1990) on the Canada lynx. They strongly recommended
that forests be managed so as to maintain good snowshoe
hare habitat-winterpatches of successional vegetation
for browse and relatively dense stands of trees for shelter.
They point out that, with a few modifications, conventional forest management is compatible with lynx habitat
management, including such practices as controlled buming, small patch (as opposed to extensive) clear-cutting,thinning stands to maximize tree growth potential, controlof pests, and construction of logging roads. In the northwestern U.S., along the southern boundary of Canada lynx
range, they also emphasize the importance of maintaining
interspersed stands of old-growth forest, which lynx in thisarea use for denning. While they note that modifications tomanagement practices, such as scheduling tree thinning
either early (when snowshoe hares have not yet recolonized an area) or late (30-40 year old growth little used byhares), may result in an initial increase in management
costs, these could be offset by greater economic and ecological diversity.

Results of (4) assessed relative to (1) and (2);Annual collection of “winter lynx abundance indices”
based on field personnel “impression”
or from fixed
winter track transects (e.g., specific vehicle, snowmobile or snowshoe routes);
Awareness by managers of all literature relative tolynx-hare cycles and an understanding of the principles that control lynx densities.
Most management programs in Canada and the U.S.
include three basic components which are discussed
below: (1) habitat evaluation and management; (2) assessments of population size, structure, and trends; and (3) harvest control and monitoring.

Habitat Evaluation and Management
The degree to which cat habitat is evaluated and managed
varies widely by jurisdiction.
In general, management
authorities (1) evaluate and categorize types and qualities
of cat habitat; (2) determine the distribution and amountof habitat in the various categories of quality; and (3)determine changes in the distribution and amount of habitat that might affect management.
The habitat use and habitat requirements of the two
North American lynxes have been extensively studied (seeSpecies Accounts), and in some cases, attempts have been
made to translate study results into workable management
tools. Habitat suitability guidelines and indexes have beendeveloped (e.g., Boyle and Fendley 1987, Anon. 1991e).
On the other hand, some jurisdictions (for example, several
western states and provinces) assume that their entire management area is comprised of lynx or bobcat habitat ofequivalent quality. Others define cat habitat on the basis ofvegetation types or physiographic regions, and base theirdetermination of habitat quality on some assessment ofcat density within each habitat type. In the U.S., for example, South Carolina has defined three categories of bobcat
habitat-coastal
plain, Piedmont, and foothill-based
upon physiogeographic regions, and has defined habitat
quality based on scent-station survey indices of bobcat
abundance within each region (B. Baker, South Carolina
Wildlife and Marine Resources Department in Zitt. to the
USFWS Office of the Scientific Authority [OSA] 1988).
Texas defines bobcat habitat as “all areas dominated bynatural vegetation,” and uses a state habitat map to determine the percentage of natural vegetation within eachcounty in each of the state’s 10 ecological regions. Bobcat
density estimates are derived for each ecological region on
the basis of a combination of harvest/population
ratiosAssessmentsof Population Size,
Structure,
and Trends
To satisfy national biological information requirements,
many provinces and states estimate the size of their cat
populations. Most have approached the problem by conducting one or more radio-tracking studies to determine
the various parameters needed to estimate population density in representative habitats (e.g., average home range
size for males and females, extent of intra- and intersexual overlap in home ranges, and proportion of transientsin the study population). The population density estimate
for each habitat type (particularly important for the bobcat,
a strong habitat generalist) can then be multiplied by the
total area for each habitat to obtain a population total.
The accuracy of these population estimates depends

230

Part II: Major

upon both the accuracy of the habitat-specific density estimates and the accuracy of the extrapolation (Rolley 1987).
Habitat-specific density estimates can be affected by thepresence of uncollared resident cats in the study area
(Rolley 1987); inaccurate estimates of home range size oroverlap due to small sample sizes; or inaccurate estimateof the proportion of transients in the population. The accuracy of the extrapolation can be affected by inaccurate
measurement of the total area of habitat in the jurisdiction
or by applying a density estimate derived from one habitat to one that is dissimilar.
Management authorities attach particular importance toreliable information on population changes, especially
population response to trapping. Some only monitor population trends, rather than estimate total population size.
Because radio-tracking studies are costly, labor-intensive,and require many years to complete, managers seldom use
them over the long term to monitor population changes.
Instead, most use indirect indices of population levels, as
described below.
The quality of population monitoring obviously varies
among jurisdictions.
Some do little more than monitor
harvest. For example, the terrestrial wildlife manager for
Colorado has reported: “While we do not have specificdata on trap effort, track indices, bobcats observed, orother good population indicators, reports from both ourfield men and the involved public indicate no noticeablechanges in population” (R.J. Tully in litt. to OSA 1989).
While a good deal of data has been collected on the
population variables described below, numerous factors
compromise their utility, and Rolley (1987) has suggested
the use of at least two indirect indices to mitigate against
uncertainty: confidence should increase if “several indicesshow the same trend.” This is borne out by a comparisonof bobcat density estimators (scent-station transects, monitoring of harvest levels, and radio-isotope feces tagging)with the number of bobcats known to be within a northeastern Florida study area over a two-year period as determined by radio-telemetry. Connor et al. (1983) found that
all indices moved in synchrony with the radiotelemetry
estimate (Conner et al. 1983). Many management authorities do in fact utilize multiple estimators. Minnesota, forexample, uses mandatory harvest reporting, mandatory
carcass collection to determine sex, age, and reproductive
parameters, annual scent-station surveys, and computermodelling of population changes (Boggess et al. 1989).

Issues.

Chapter

4. Trade

cyclical highs of Canada lynx harvest-fromnearly80,000 pelts per year in the late 1880s to about 20,000 30
years later, with a continued decline through 1940 (Eltonand Keith 1946: see Fig. 6 in the Canada lynx Species
Account). The period 1880-1920 was a time when western Canada was opening up, trapping was essentiallyuncontrolled, and there was intense competition for fursamong unregulated trappers. Harvests only began toclimb again in the 1960s possibly reflecting an increase inlynx numbers following institution of management programs (Todd 1985).
However, change in harvest levels can be influenced byfactors other than the size of the cat population. As Rolley(1987) has written: “Annual harvest estimates will reflectchanges in population levels only if harvest pressure isrelatively constant.” Harvest pressure, however, is rarely“relatively constant,” being influenced strongly by winterweather (Rolley 1987), number of trappers (Tewes andScott 1987), and pelt prices (Erickson and Sampson 1978,
Brand and Keith 1979)-harvest
tends to increase withpelt price, although some evidence is contradictory (Govt.
of Canada 1988).
Data on harvest effort collected by trapper interviewnumber of cats caught per trapper, number of trap-nights
per cat, or number of cats per trap-night (Slough and Ward
1990, Tewes and Scott 1987, R. Eagan, pers. comm.)can be a better index to population change because biases
related to pelt price and total number of trappers are eliminated. The principal concern is the accuracy of the harvest effort measurement.
Age and Sex Ratio Changes
Patterns of, and changes in, age and/or sex ratios are often
used as indices of population change. This is particularly
true for cyclic Canada lynx populations. In Canada, mostauthorities routinely measure pelt size (tip of nose to baseof tail) prior to auction to distinguish adults from youngof the year, or kits (Quinn and Gardner 1984, Slough andWard 1990, Poole 1992). Monitored on an annual basis,a peak in the proportion of kits in the harvest should precede a cyclic decrease in lynx productivity and a fall in
the population size. However, time of collection can influence kit representation within the overall sample because
the number of kits caught increases with the length of thetrapping season (Parker et al. 1983, Quinn and Thompson
1985, Quinn and Parker 1987).
Some authorities collect carcasses from trappers.
These carcasses can be sexed, and age can be determinedby sectioning teeth and counting the rings of cementum
annuli (Quinn and Parker 1987, Rolley 1987). Age-specific fecundity and mortality rates can be determined.
Several studies of bobcat and Canada lynx populations
have pointed towards vulnerability to trapping which isboth age- (yearlings may be more vulnerable to trapping as

Harvest Level and Harvest Effort
Changes in overall harvest or harvest effort (measured ascatch per unit of harvest effort) can be used to help evaluate population status. Change in overall harvest is a crude,
but still useful, indicator of abundance. For example,
analysis of fur harvest records kept by Canada’s Hudson
Bay Company show a striking difference over time in the231Part II: Major

issues.

Chapter

4. Tradecomputer models to determine the effects of various harvest rates on population parameters, including population
size. An acceptable harvest rate is then selected-a rate
which maintains the population at the pre-determined optimal level and allows for environmental fluctuations which
may increase natural mortality or decrease recruitment andsurvival. Other authorities resort to “rule-of-thumb,” “bestguess estimates,” or tradition to set harvest levels. For
example, in the U.S., a year’s quota is often set at the samelevel as the previous year’s, especially if this level of harvest has been sustained over several years without adecrease in population indices. For the Canada lynx,
annual quotas are set with regard to previous year’s harvest
and the current position of local populations within thelynx-hare cycle (Quinn and Parker 1987, Rolley 1987). It
has been argued that it is not necessary to estimate population size prior to harvest commencement, but rather thatharvest level can be adjusted according to population
change indices (e.g., Slough and Jessup 1994).
While many jurisdictions calculate “safe” harvest levels, not all institutionalize these calculations as maximum
allowable quotas. Canada lynx harvest is regulated byannual quota in Manitoba and parts of Ontario and Alberta(Canada), and Montana and Idaho (U.S.). Bobcat harvestis regulated by annual quota in Arkansas, California,Louisiana, Massachusetts, Minnesota, Montana, Oklahoma, Oregon, South Carolina, Tennessee, Virginia, West
Virginia, and Wisconsin (U.S.), but nowhere in Canada.
Quotas may take the form of total animals allowed caught
per jurisdiction (i.e., 500 bobcats per year in Louisiana)or total animals allowed caught per person (i.e., in Ontarioannual quotas are established per registered trapline
through the cooperation of local wildlife officials and thetrapper) (Govt. of Canada 1983, 1988; Govt. of U.S.
1983a,b; Johnson 1990, Koehler 1990).
Other forms of harvest regulation include manipulation of harvest season length and chronology, area of harvest, take methods, and licensing requirements.
New
regulations are promulgated from time to time, and areusually published well in advance of the open season. All
jurisdictions set a season length for harvest of bobcat andCanada lynx (both for hunting and trapping), generally in
winter (November to February), so as to avoid taking
breeding adults and immatures. If harvests are felt to havebeen too high in recent years, management authorities may
close their season indefinitely, either in the whole or part oftheir jurisdiction. Some jurisdictions prohibit trapping incertain areas which serve as refugia from which cats candisperse to recolonize hunted areas. Licensing requirements vary widely, from the simple purchase of a season
license in some U.S. states to more complex legal systemsof long-term leasing (five-year terms in the Yukon) of registered trapping concessions in some Canadian provinces(Govt. of Canada 1983,1988; Govt. of U.S. 1983a,b; Todda result of their dispersal over unfamiliar areas) and sexspecific (males may be more vulnerable to trapping
because their home ranges are larger and thus contain more
traps) (for summary see Anderson 1987, Quinn and Parker
1987, Rolley 1987). Some models for estimating population size from the harvest have attempted to take such vulnerability into account (e.g., Paloheimo and Fraser 1981).
However, disparities in age and sex ratios in the harvest
must be weighed against the fact that age and sex ratios
within the population can be strongly affected by density(Frederickson and Rice 1979, Lembeck and Gould 1979,
Zezulak and Schwab 1979, Slough and Ward 1990). For
example, more males than females in the harvest could
reflect the presence of more males in the population, rather
than simply greater male vulnerability to trapping.
Changes in Prey Availability
As discussed in the species account, throughout much of
their range lynx populations fluctuate cyclically along with
their main prey, the snowshoe hare, typically lagging one
to two years behind. Several Canada authorities are researching the effectiveness of monitoring snowshoe hareabundance (through a combination of feces counts, track
counts, and trapper interviews) as an index of lynx population change (Slough and Ward 1990, Poole 1992).
Changes in prey availability are also a pertinent index
to bobcat populations in some areas. For example, Knick(1990) studied bobcats in Idaho at a time when lagomorph
populations were declining, and found that bobcat numbers also declined.
Scent-station Surveys and Track Counts
These two methods of estimating population size, and theirstrengths and weaknesses, are discussed in the Researchchapter.
Population ModelingManagement authorities have developed a variety of computer models to estimate changes in bobcat and Canada
lynx populations which utilize field data collected by theabove methods on population levels, productivity/recruitment, population age and sex ratios, and harvest and nonharvest mortality rates.

Harvest Control and Monitoring
Management of bobcat and Canada lynx harvest involves,for the most part, determination of allowable harvest levels
and control of the harvest so that it does not exceed theselevels. Management assumes that cat populations are atsome equilibrium level below carrying capacity, and therefore produce a harvestable surplus.
Determining allowable harvest levels, or quotas, is animprecise exercise. Some authorities use sophisticated232Part II: Major

Chapter

4. Trade

Anderson 1987).
Similarly, Canada lynx pelt prices increased over thesame period. During the cyclic low in lynx populations
in Canada in the mid- 1970s provincial harvests were up to
three times higher than during the preceding two cycles.
However, during the peak in lynx abundance in the late1970s-early 1980s provincial harvest levels were 40-70%
lower than during the preceding peak (with the exceptionof the Yukon and Northwest Territories, where the peakharvests were slightly higher, although this probably
relates more to increased trapping pressure than to actual
population increases: Todd 1985, Govt. of Canada 1988).
This was a reversal of the apparent recovery trend of
Canada lynx since the early 1960s. A similar trend, on asmaller scale, has been documented in the American statesof Washington and Montana (S. Thorpe in Zitt. to USFWS
199 1). Moreover, the highest prices historically recorded
for lynx pelts (up to U.S. $750 per pelt) were offered in themid- 1980s when lynx populations again were at their
cyclic low, and harvests again were high (Govt. of Canada
1988).
Todd (1985) considers various factors that could
account for the reduced peak lynx harvest, including habitat loss, severe winter weather, and declines in snowshoe
hare numbers, but concludes that the most likely explanation is overtrapping during the cyclic low. At this time,overharvest would theoretically reduce the breeding population necessary to regenerate a peak, and thus produce anoverall decline in lynx abundance (Brand and Keith 1979).
Shortening or closure of harvest during cyclic lows has
been called for (Berrie 1973, Brand and Keith 1979,
Carbyn and Patriquin 1983, Parker et a2. 1983, Todd 1985,
Bailey et al. 1986, R. Eagan, pers. comm.), and this recommendation has been implemented in parts of somestates and provinces (Govt. of Canada 1988). However,
Poole ( 1994) believes that stopping trapping during periods of low hare densities in the Northwest Territories is notnecessary. His evidence suggests that trapping during atleast the first two winters of low hare densities may bepartly compensatory, as trapping of lynx likely to starve
would have little impact on the population. In addition, asignificant portion of the entire community trapping area
appears to be buffered from trapping and may provide substantial refuges for population recovery.
An alternative management strategy involves maintenance of refugia in good-quality habitat to ensure protection of breeding lynx nuclei during cyclic lows. This
strategy would appear more appropriate for areas where
there is no evidence for significant lynx population
declines, such as north-western Canada. Studies to identify and define such refugia, and investigate their role inlynx trapping areas, are ongoing in the Yukon andNorthwest Territories (Slough and Ward 1990, Poole
1994). Poole ( 1994), for example, has found that a signif-

1985, Slough and Jessup 1994).
Harvest monitoring is‘accomplished by a variety ofmethods. Both the U.S. and Canada require that pelts destined for international export be tagged with nationallyapproved tags; many jurisdictions require that all pelts betagged. Some jurisdictions require that all carcasses besubmitted for examination of age, sex, and reproduction; inothers, this is voluntary. Some management authorities
require certain information from trappers, hunters, or both(e.g. Idaho requires all licensed trappers to prepare anannual report); others require registered fur dealers toreport.

Are Canada
Sustainable

Issues.

Lynx and Bobcat Harvests
as Presently Managed?

IUCN has recently drafted a set of guidelines to help
define when the use of a wild species is sustainable (IUCN
in prep). According to their criteria, a use of a wild species
is likely to be sustainable if:
1. It does not reduce the future use potential of the target
population or impair its long- term viability;
2. It is compatible with maintenance of the long-term viability of supporting and dependent ecosystems;
3. It does not reduce the future use potential or impair the
long-term viability of other species.
Since rare species are seldom trapped incidentally to
Canada lynx or bobcat, and since habitat management forthese cats (where practiced) is compatible with conventional forest management, the last two criteria are satisfied.
With regard to the first criterion, the American andCanadian governments have invested substantial effort inensuring that harvest levels do not reduce the viability of
their bobcat and lynx populations. Annual reported harvests for bobcat and Canada lynx over the past 15 years arepresented in Figures 4 and 5. For the bobcat, the primary
period of concern was the late 1970s to early 1980s. Prior
to this, bobcat pelts had little value (the average pelt pricefrom 1950- 1970 being only U.S. $5- 10: Young 1958) andbobcats were generally considered pests. Harvest levels
greatly increased following the entry into force of theAmerican Endangered Species Act in 1969 and CITES in
1975, when the pelts of cats listed on Appendix I werebarred from legal trade, and the value of bobcat pelts
increased to hundreds of dollars (see Species Account).
The U.S. government temporarily banned export of bobcat
pelts in 1977 pending evidence to indicate that harvest levels were not detrimental to the survival of the species,
prompting state governments to develop and institute harvest management programs, and leading to a huge increasein field research on the species (Blum and Escherich 1979,

233

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500

Figure 4. Annual harvest
and averagepelt price forthe lynx in Canada.
Source: Canadian
Wildlife
Service, Statistics Canada,
and TRAFFIC
U.S.A.

400

300

200

10 0

0

Figure 5. Annual
harvestand averagepelt pricefor the bobcat
in theU.S. Source: U.S. Fish
and Wildlife Service and
TRAFFICU.S.A.

234

Pat? II: Major

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4. Trade

could play an important role in maintaining harvested
bobcat populations, as well.

icant portion of his community trapping study area in the
Northwest Territory is apparently buffered from trapping,
due to natural features and other factors, and thus provides
a substantial refuge for population recovery.
Management of North American lynx and bobcat harvest has thus been adaptive-both harvest and populationlevels are monitored, and regulation of exploitation ismade more restrictive if evidence arises to indicate that use
is potentially unsustainable. Research has been an integral
part of resource management, and management and population/harvest monitoring techniques have become progressively more refined and sophisticated. This is one of
the requirements defined by IUCN (in prep.) for management for sustainable use: “A management system isneeded that is able to adapt and adjust uses in response tochanges in the target population, its supporting ecosystems, and other affected species. Such a system recognizes
that all the information needed to ensure sustainable use
may not be available. It therefore sets use levels cautiously
and adjusts them in response to monitoring and othersources of information.”
While biologicalknowledge
and management
resources for the Canada lynx and bobcat far outstrip thoseof any other cat-countries in the world, they are still not
sufficient to yield the information needed to guarantee thatharvests are sustainable-reliable
periodic population estimates. Given the difficulties of counting cats (see Chapter
3, Research), however, such estimates are likely to remain
unavailable for some time. Yet it is probably safe to saythat current North American management practices haveresulted in sustainable harvests in that they have been sufficient to prevent widespread and prolonged overharvest.
Under such a management regime, the long-term viabilityof both cat species is unlikely to be impaired, and the commercial use of bobcats and Canada lynx can thus be considered sustainable. Additional system refinements areneeded for better insurance against local population depletion, however, such as appears to have occurred in somesouthern populations of Canada lynx.
A number of suggestions have been put forward for
improvement of management practices (Brand and Keith
1979, Todd 1985, Bailey et al. 1986, Quinn and Parker
1987, Rolley 1987, Govt. of Canada 1988, Knick 1990,
Tewes and Scott 1987). These authors acknowledge thatharvest regulation is a jurisdictional responsibility, butnonetheless recommend a certain degree of standardization, including the following:

All harvests should be regulated at the appropriate level(trap-line, state- or province-wide) and established byanalysis of biological and survey data. However, quotas may not be necessary for cats in remote areas wheretrapping pressure is more dispersed (or, for the Canada
lynx, at least not during years of high recruitment).
Harvest monitoring procedures should be standardized.
It has been suggested that (a) all jurisdictions shouldcollect age/sex data from pelt measurements and carcasses by standardized procedure to gauge the statusof the population and, in the case of the Canada lynx, its
position in the hare-lynx cycle; (b) all jurisdictions
monitor trapping effort and trapping success. Tewes
and Scott (1987) suggest that the responsibility formonitoring harvest in the U.S. should be decentralized
to the county or management unit level.
Harvest reporting should be standardized, particularlyin the U.S.
All pelts should be tagged or sealed within the jurisdiction of harvest, and taxes/royalty rates should bestandardized to discourage illegal inter-jurisdictional
movement of pelts for the purpose of avoiding higher
taritrs.
If a use is sustainable and long-term population viability
is not affected, the question of whether the use is conducive to species conservation is not relevant. However,
not only is there sufficient room for doubt regarding thebiological impact of past harvests on numbers of Canada
lynx and bobcat but, at present, world demand for thesecats’ fur is declining (Table 1). Under such circumstances,
it is important to ask whether commercial use can be considered to support the conservation of these species.

Does Commercial
Use Benefit Bobcat
and Canada Lynx Conservation?
Fur trapping entails removal of animals from wild populations and can benefit these populations if it serves as anincentive to relieve other population-reducing pressures,primarily habitat loss. IUCN (in prep.) recognizes that“the social and economic benefits from sustainable use
could provide powerful incentives to conserve wild
species and their supporting ecosystems, providing that (1)the people most likely to have a direct impact on thespecies and ecosystems concerned receive a fair share of
the benefits from the use; and (2) there is a clear connection between the benefits and conservation” (e.g., “a portion of these benefits should be reinvested in maintaining

1. There should be simultaneous application of harvest
strategies over broad eco-regions.
2. In the case of the Canada lynx, trapping seasons should
be closed or shortened during cyclic population lows, ora protected refugia strategy should be implemented.
Knick’s (1990) computer models indicate that refugia

235

Part II: Major

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Chapter

4. Tradedrug trade is a good example. It is unlikely that banning
import of spotted cat pelts into Europe would have been so
successful had it not been for a major decline in demand.
Illegal wildlife trade remains a serious problem even for
the United States, where CITES issues are accorded relatively high priority by the government. The U.S. Fish andWildlife Service has estimated that for every 10 wildlife
shipments entering the country legally, one to four enter
undeclared and undetected (Grove 1981). In 1988, 65
inspectors were faced with the task of inspecting more than
83,000 declared shipments of wildlife for compliance witha number of national regulations as well as CITES regulations. In reality, the inspectors only managed to inspect
about 25 percent of the declared shipments (Mulliken andThomsen 1990).
The significance of illegal trade in cat products isshown by the following example. In 1990, the majorityof exports of tiger bone medicines reported by China wereto the United States. Although the United States prohibits
import of tiger bone, neither import nor seizure wasreported in 1990 (Headley 1992).
Illegal trade in cats and their products generally
assumes three forms: trade in live animals as pets, trade inpelts or body parts (claws, teeth, etc.) for fur garments ornovelties, and trade in bones and other body parts for
Oriental medicine. Of the three, the latter is probably themost serious. Demand for the first two categories is probably relatively low and unlikely to be increasing. Tourists
make up an important proportion of consumers.
It is more difficult to evaluate demand for bone fromthe medicinal trade because, as will be discussed below, abasic understanding of the dynamics of the consumer market is lacking. Demand may be increasing given the recent
reports of tiger poaching from India and Nepal, but it may
also be decreasing or unchanged-thepoaching in the
Indian subcontinent does not necessarily indicate a change
in consumer demand, but could reflect widespread depletion of supply. In other tropical Asian tiger range states,
tiger conservation is not of such high priority and poachingincidents are less likely to gain media attention. The statusof the tiger in these countries is highly uncertain.
Medicines containing tiger bone or other cat bones areused primarily by Chinese and Koreans. Consumers are
not limited to these two countries, however, but are foundthroughout the world, mainly where there are Oriental
communities.
Of all the threats facing cats, illegal trade has the greatest potential to do maximum harm in minimum time.
Commercial poaching can be devastating to species-witness the well-documented
cases of the black rhino andAfrican elephant. The existence of commercial poaching
and illegal trade indicates product demand, but when the
product is illegal, it can be very difficult to learn enoughabout the nature of the demand to take appropriate correc-

target populations and their supporting ecosystems”).
The furs of the bobcat and Canada lynx are among themost valuable of North American furbearers (Shieff and
Baker 1987). When prices were at their highest in the mid1980s the trade in Canada lynx pelts provided some
U.S.$3,800,000 directly to trappers (Canadian Wildlife
Service in litt. 1994). As Todd (1985) pointed out, the capture of even one Canada lynx represented a significant
contribution to a trapper’s income. Even for the bobcat,
or for the lynx in the current low-price market, cat fur revenues are still relatively significant. In the 1987- 1988
California trapping season, for example, the average pertrapper income from an average harvest of 13.5 bobcats
came to just under U.S.$2,000 (Gould 1989). Do such revenues serve as an incentive for trappers to actively support cat conservation?
The answer to this question is equivocal. Looked at one
way, the answer is “no.” The highest pelt prices offered
for Canada lynx were at a time when the population wasat its cyclic low, a basic function of supply and demand,and there is justifiable concern that over-trapping has
reduced populations. People often place short-term gain
over long-term interest: in the words of Todd ( 1985),“even the most conscientious registered trapper may be
tempted to over-trap lynx when they are valuable, despite
hoping for the sake of the lynx that the price will go down
soon .” This is why government regulation is necessary to
avoid overharvest.
On the other hand, hunters and trappers form an important powerful constituency for conservation of wildlands inboth Canada and the U.S. License revenues and other tariffs derived from cat hunting also contribute to the budgets of government agencies charged with ecosystemconservation.
There have been no studies examining to what degree
either consumptive or nonconsumptive use values of catstranslate into effective cat conservation. As with the valueof sport-hunted cats, the commercial value of North
American cat pelts is unlikely to be a major force for cat
conservation if viewed in isolation from other values,
including the existence value many people place on wildcats, as well as other wildland-derived economic revenues.
However, those who derive commercial benefit from theharvest of North American cats are among the groupwhich lobbies for actions contributing to the conservationof these species.

Illegal Trade in Cat Products
Illegal trade takes place when legal trade is restricted or
prohibited but some degree of consumer demand remains.
When that demand is strong, it is very difficult to prevent
trade through legal controls alone-the resilience of the236Part II: Major!-iger and leopard+wx seized from illegal traders

Issues.

Chapter

4. Tradein Calcutta in 1989.

)es not necessxily
ow th at the\: will reduc’c’ ortrary , blXk nxuket
denxllId--on the‘31\stiyutf’ could actually increase wnsunw
intert’st if tr;tdc
bms are not strictly
Oriental medicine trade is one
posed to cat conservation today.
three

the
prottxti
and the tr;lde i5il

Megal Trade in Pelts and Live Animals

Part II: Major

Issues.

Chapter

4. Trade

Skins displayedin Tachilek

on Myanmar

frontierwith Thailand

in 1994.

1993, but prior to that it was nonetheless required, throughmembership in the European Union, to enforce EU CITES
regulations. TRAFFIC Europe (de Meulenaer and Gray
1992) surveyed fur shops in several major Greek cities, andfound coats made from Appendix I cats in over 100 ofthem. Many garments were made from small spotted fur
scraps, but others were made from full pelts. Species intrade included ocelot, leopard, cheetah, and jaguar. While
TRAFFIC states that some of these skins may have been
imported legally, they were concerned about the lack ofmonitoring by Greek authorities. Traders did not help toallay suspicions by proposing several ways of avoiding
controls (de Meulenaer and Gray 1992).
The consumer market consists largely of tourists, ofwhich Greece is host to some 10,000,000 annually.
TRAFFIC notes that this is particularly worrisome for EUwide enforcement of CITES: with internal border controls
being removed, each member state becomes a potential
door to the entire EU market.
In Asia, Kashmir has long been a center of fur processing (van den Berg and Damhuis 1982, Verma 1983,
Chopra 1988, Mohapatra 1988). Coats made in Kashmirfrom snow leopard, leopard, clouded leopard, fishing cat,leopard cat, jungle cat, wildcat, and rusty-spotted cat havebeen found for sale recently to tourists in Kathmandu,
Nepal, in a Kashmiri-run fur district (Barnes 1989, vanGruisen and Sinclair 1992). As in Greece, traders provided advice on how to avoid western import controls. In
China, the fur tanning industry in the northeastern city ofHarbin is similar to that in Kashmir, and well processedskins of leopard and Asiatic golden cat have been foundis little evidence in the former consuming countries ofEurope, North America, and east Asia of significant
demand for garments made from species which were once
popular, and are now protected (such as the leopard or
ocelot). However, there is still evidence of illegal trade inpelts in many of the range states with spotted cats. In Latin
America, large shipments of pelts are occasionally confiscated. For example, 675 pelts, mostly oncilla, were seized
en route from Paraguay by Brazilian authorities, and wereincinerated in February 199 1 in Iguacu National Park (P.
Crawshaw, A. Ximenez in Zitt. 199 I).
Montevideo (Uruguay) and Buenos Aires (Argentina)
were major centers of the illegal fur trade in Latin America
up until the early 1990s. The main felid furs in trade wereobtained from Geoffroy’s and pampas cats. Since 1990,
there has been a significant improvement in the enforcement of international trade controls in both Argentina and
Uruguay. Illegal trade in the furs of these species has
decreased dramatically as a result. However, very smallinternal markets for fur items persist in Montevideo andBuenos Aires, in spite of legislation banning the sale offurs of these species. Dealers involved in the illegal trade
employ various tactics to make the garments they sell
appear old, and therefore not subject to government sales
controls. One method involves sewing labels from old
European fur garments into garments offered for sale;
another involves invoicing buyers of fur coats for a“repair” rather than a “purchase” (J. Villalba-Macias. inlift. 1993; Tomas Waller, pers. comm. 1994).
In Europe, open sale of coats made from cat furs appears
to be restricted to Greece. Greece did not join CITES until

238

Part II: Major

Issues.

Chapter

4. Tradethe roof, its nose will induce the birth of boys. Officials
wore the “majestic bone” (or clavicle) of the tiger aroundtheir necks to give them poise and inspire the envy of thecommon people. Tiger whiskers cure toothache. There
are many uses for tiger bones, with curative powers for adiverse range of complaints from rat-bite sores tohydrophobia, but chiefly having to do with building strong
bones and teeth. Sliced and ashed, bones taken from tiger
feces cure alcoholism-in
November 1992, the tiger
enclosure of the Taipei Zoo saw an upsurge of demand
from wives anxious to end their husband’s chronic dririking (K. Nowell, pers. obs.). Tiger penis makes tigers ofmen (Read 1982, Martin 1987).
Modern Chinese pharmaceutical texts do not dwell onsuch claims, and discuss only tiger bone, prescribed
mainly to alleviate symptoms of rheumatic and bone marrow disease (Anon. 1976). Tiger bone is often combinedwith other ingredients to make a plaster for aching joints
and sore muscles. Small portions of ground tiger bone
are mixed with liquor to make tiger bone wine, which ismore of an invigorating tonic, with aphrodisiacal overtones, than a medicine.
Tiger bone is marketed in two ways. Traditionally,bones are stocked by pharmacies and doctors, and small
portions are sliced off and ground on the spot for patients.
Leg bones are favored, particularly the humerus-proba-

for sale there (Low 1991).
Skins of protected cats ‘and live wild-trapped animals,
both typically in poor condition, can be spotted in wildlife
markets around the world. This trade, while illegal, isoften highly visible, especially to visiting conservationminded westerners, and the Cat Specialist Group hasreceived numerous reports from dozens of countries, richand poor. Because hunting intensity varies locally, it isdifficult to draw any conclusions about the overall impactof this trade on cat populations. In general, however, thesemarkets are more likely to reflect local levels of persecution of cats (for livestock protection, sport, opportunity,occasional income, etc.), rather than systematic commercial poaching of the sort that is now being directed at thetiger, as discussed below.

The Bones of a Dilemma:
Tigers and Oriental Medicine
Ancient Chinese medical texts dating back some 2,000
years proclaim that tiger flesh improves vitality and is atalisman against 36 kinds of demon. Tiger fat is a curefor hemmorrhoids. Tiger blood builds up the constitution
and strengthens the willpower. Tiger testes treat scrofula.
Tiger eyes clarify the vision and stop crying. Hung from

Two tiger humerus
bones at right show foramina(holes for nerves and blood vessels) at the distal
ends, which are distinctiveof big cats. Third from the right is a femur bone. At left, the smaller
humerus
may be from a leopard, a species also used in traditional
Oriental medicine.

239

Part II: Major

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Chapter

4. Trade

bly not only because these are the more “powerful” preycatching limbs, but because the humerus has a large foramen (an opening allowing the passage of blood vessels andnerves) at its distal end, distinctive to the family Felidae,
which permits the authentic to be distinguished from the
fake. In Taiwan, wholesale prices for such bones ranged
between U.S. $860-1,280 per kg (about five humeri) in
October 1992 (Nowell 1993a, b). Traditional Oriental
pharmacies, which stock a variety of herbs and animal
parts, are found throughout east Asia and in overseas east
Asian communities throughout the world, which number
some 29,000,OOO people (Anon. 1992k).
The second, more modern form, is in manufactured
medicines, including pills, powders, and wine. Up until
1994, China and South Korea were the chief producers.
China’s annual reports to CITES over the 1980s and early
1990s indicate exports to Hong Kong, Malaysia, Singapore,
Thailand, and the United States. Hong Kong, which plays
a key middleman role in China’s traditional medicine trade,is consistently reported as the major exporter of tiger products seized in the U.S. (Headley 1992).
However, in the face of threats of trade sanctions fromCITES Parties and from the United States, China
announced in May 1993 that it would ban production oftiger bone medicines, at that time manufactured by more

Tiger specialists

than 20 companies in over 100 factories (Tan 1987, X.
Liu in Zitt. 1992), following a six-month grace period. The
South Korean government also announced prohibition ofdomestic trade in tiger bone in May 1994, with a ban on
production to take effect in November, and a ban on sale oftiger bone products to follow six months later. They
claimed that all tiger bone stocks in the country had been
identified and registered as of March 1994 and individually marked by the government in May (J. Thomsen in
Zitt. June 1994).
Although nearly every Asian range state has protected its
tigers from hunting and banned international and internal
trade in their products, medicines labelled as tiger bone
have been found for sale throughout the region (Tan 1987,
Martin 1992a,b, Anon. 1992i), as well as in many westerncountries. Poaching is widespread, commercialized, anddriven by the bone trade. For example, in India, tiger carcasses have been found with the bones removed and theskins left behind (S. Dey, pers. comm. 1992). According
to C. McDougal (in Zitt. 1991), bones are sold in Bangkok at
prices 10 times higher per kg than pelts. The threat goesbeyond the tiger to encompass other Asian cats: the bonesof leopard, snow leopard, clouded leopard, golden cat, andleopard cat are sold as substitutes for tiger bone (Tan 1987,
J. Yu in Zitt. 199 1, R. Jackson, pers. comm.).

in 1995 examining

240

canine

teeth

made

into pendants

in a Hanoi

curio shop.

Part II: Major

Issues.

Chapter

4. Trade

Many fake tiger parts are offered for sale, such as these penises
at Tachilek in Myanmar(left). Above, the genuine article isdepicted in a Chinese publication(Lin, H. and Chen, S. 1988).

macies where the proprietors were clearly taken aback tolearn that they were fakes (K. Nowell, pers. obs.). The
price of an item marketed as a tiger derivative is not necessarily an indication of its legitimacy, but rather of theseller’s audacity.
Moreover, the Wildlife Forensics Laboratory of the
U.S. Fish and Wildlife Service has tested more than 10
samples of Chinese-manufactured“tiger bone” plasters
and powders, and found that the products actually contained no bone of any kind (E. Espinoza, pers. comm.
1993). This finding not only complicates law enforcement, but points out how little is understood of the tiger
bone trade. It is not at all clear how much tiger bone-orother wild cat bone-is
actually being used to produce
manufactured medicines. Chinese industry sources havereported that it takes 250 g of tiger bone (roughly theweight of a humerus) to manufacture 55 kg of tiger bonewine (Tan 1987). While no official import statistics for“raw tiger bone” are available from China, customs datafrom South Korea shows average annual imports of 340 kg
between 1975 and 1992, at an average price of U.S. $127
per kg (J. Mills 1993). According to traders in southeast
Asia, the yield of dried bone from a single tiger ranges
from 6- 11 kg (Anon. 1992i), which implies that South
Korea imported the equivalent of 557- 1,021 tigers over
18 years (if the bones were really tiger bone).
However, with the supply of wild tigers declining, perhaps real tiger bone is being diverted to the more traditional pharmacies, where discriminationof fakes issomewhat easier. In October 1992, real tiger bone wasbeing sold in the main wholesale district in Taipei, Taiwan
in October 1992 (Nowell 1993a, b). Following publicity
campaigns launched by several western environmentalgroups, the Prime Minister held a press conference to reiterate that trade in tiger bone has been banned since 1989,
and called for better law enforcement.
According to

Unfortunately, the tiger bone trade has not been systematically investigated, and its organization and dynamics are largely a puzzle. For example, sources in India and
Nepal identified China as the destination for tigers
poached in these countries (Martin 1992c, Anon. 19921).
The medicinal manufacturers would seem to be the obvious culprit. On the one hand, the Chinese government has
claimed that all stocks of tiger bone used by the manufacturers were obtained prior to 198 1, when China joined
CITES, which permits legal export of their medicines as“pre-Convention.”On the other hand, the government
has confessed that it cannot “calculate exactly how muchtiger bone is used in our country at present,” nor “exactly
how many factories are producing medicines which
include tiger bone” (X. Liu in Zitt. 1992). However, it isobvious that new stocks of tiger bone have entered thecountry since 198 1. Smuggling of tiger parts from Burma
to China was reported from as early as 1983 (Salter 1983,
Anon. 1984), and Tan (1987) reported that the medical
industry was paying what was then considered a very highprice of U.S. $200 per kg for Burmese tiger bone.
Yet reaching simple conclusions about the ultimate destination of poached tiger bone is difficult because what issold as “tiger bone” is often not genuine. Oriental medicine has its skilled practitioners, but also its share of charlatans. Faking valuable natural medicinal ingredients issomething of an art fort-n, and the Chinese have devoted afull color photo textbook to distinguishing the false fromthe real (Lin and Chen 1988). The book’s photos clearlyshow that products which are marketed as tiger parts (andhave been reported as such by investigators) are actuallytaken from cattle. This includes “tiger” penis and “tiger
legs”; the latter can be easily found in China sold by“Tibetan” street peddlers, and are put together from cattlebone and ligaments, a bit of striped orange fur, and dog
claws. They have been seen in Taiwanese traditional phar-241

Part II: Major

Tiger

bones

Issues.

seizedChapter

4. Tradethat such products were not for sale in their shops.
Legislation to increase penalties for illegal trade in wildlife
products was sent to the Legislative Yuan. The American
government recognized Taiwan’ s progress in curbing illegal trade by dropping the Pelly sanctions a year later, inJuly 1995.
The tiger poaching situation has been likened to theearly days of rhino poaching, when no one realized how
severe and thorough a commercially driven poaching crisis lay ahead (Jackson 199 lb). In the case of rhino horn,
trade bans led to international trade being taken over bycriminals and speculators. They inflated the price of thehorn, and the added cost was passed on to patients by thedoctors and pharmacists. (The 1980s were a period of significant economic growth for Asia, the main consuming
market.) Publicity regarding the endangered status of therhinos appeared to influence speculator interest in acquiring horn “before it’s too late,” which further fueled poaching. This faceless but powerful segment of the market may
actually be its driving force-investingin the rhino’s
extinction, thus inflating demand above actual levels of
consumption, which may be quite low. Given that rhino
horn does not have to be displayed in order to be prescribed, law enforcement is difficult. Meanwhile, while
the Asian medical community is generally aware of therhino’s endangered status, it stubbornly defends the useof its horn as an important medicine for serious illnesses
only (Nowell et al. 1992, Milliken et al. 1993).
To what extent does the market for tiger bone resemble that for rhino horn? On the one hand, unlike rhino
horn, tiger bone is not considered a potentially life-saving
medicine. However, like rhino horn, genuine tiger bone
is a high-priced, “charismatic,” long-lasting commodity
suitable for investment by speculators, if they believe themarket will persist despite its illegality.
At present, the internationally favored solution to crisesof commercial poaching has been to ban the trade, and toattempt to reduce demand through publicity campaigns.
This approach was successful for the African elephant andfor the spotted cats of the southern hemisphere. Its effectiveness has not yet been demonstrated for species usedby the Oriental medical trade, which is an entirely different
sort of market. Trade bans alone are unlikely to be effective if demand persists.
How strong is the demand for tiger bone? What sortof people are the main consumers, and why do they take it?
How would they react to (a) a campaign emphasizing thetiger’s endangerment; (b) a campaign to promote substitutes for tiger bone (if indeed there are any); or (c) news
that manufactured tiger bone medicines are fake? The
point is that a better understanding of consumer demand
and the marketing channels is needed before an effective
strategy for change can be developed. It is not enoughin Delhi in 1993.

TRAFFIC Taipei, all bone of any sort soon disappeared
from shop windows (J. Lob, pers. comm. May 1993).
However, deciding that Taiwan had still not effectively
closed down its internal market, the U.S. government
announced in April 1994 that it would enact limited trade
sanctions against Taiwan. as authorized by the so-called“Pelly Amendment” to the Fishermen’s Protection Act of
1967, which provides for the President to prohibit importation into the U.S. of wildlife products from an offending
country if advised that the nationals of that country are
engaging in trade or taking which diminishes the effectiveness of any international program for endangered or
threatened species. The Taiwan government responded byestablishing a special police task force, something long242

Part II: Major

Issues.

Chapter

4. Tradeant dilemma: whether or not to breed tigers for their bones
in order to save the remnant wild population.

that the tiger’s rarity has been the focus of media attention(Anon. 19921, Browne 1992, Hamer 1992, Anon. 19931,
Hennessey 1993, S. Mills 1993, Tiger Trust 1993, Linden
1994), and that most major consuming nations are rushing to demonstrate their goodwill by banning the trade.
Indeed, there is the danger that publicity and “black market
mystique” could make the problem worse, as for the rhino.
Although the Chinese government banned trade andproduction of tiger bone medicines in 1993, in 1994 it
revived a request made to CITES in 1992 for recognitionof the first commercial captive breeding operation for the
Amur tiger. Recognition would allow it to market tiger
products derived from captive tigers internationally. The
proposal, to have been submitted to the ninth Conferenceof the Parties to CITES in November 1994, involved saleof bones and other derivatives obtained from tigers whichdied or were culled at a breeding center at Hengdaohezhi
in Heilongjiang Province. The proposal was again withdrawn, this time before being officially circulated to theParties. Because the issue keeps resurfacing, it is worth abrief discussion.
The Chinese proposal stated that tiger parts would becumulatively obtained from “culled and dead individuals”
and is ambiguous about how they would be sold, but thepremise is that “limited legal trade of the derivatives andproducts of the tiger will reduce the poachers’ desire on thewild population” (Govt. of China 1994). Recent articles in
Asiaweek (Anon. 19931) and Time (Linden 1994) magazines have mentioned the idea of supplying the tiger bonemarket from captive animals. A glance at the subsequent
letters to the editors showed that while westerners are generally outraged by the idea, east Asians, particularly
Chinese, are more supportive.
Tiger farms exist in
Thailand with openly commercial aims: one owner statedrecently that, “Our scientific research will look into thepossibility of using tigers as farm animals, breeding them
the way we breed pigs” (Anon. 19946). In fact, it is likelythat there are more tigers in captivity (including “unofficial” institutions) than in the wild, and tigers are breeding
so well in recognized zoos (see Chapter 5) that “surplus”
tigers are not infrequently euthanized. If the recent tradebans and proposed consumer education campaigns do not
alleviate the tiger poaching crisis, so that the decline
towards extinction of the tiger in the wild continues, theconservation community may be faced with an unpleas-

Summary

and Conclusions

1. The fur trade is not a serious threat to spotted cats forthe foreseeable future. The only species that can be said
to be threatened by potentially unsustainable exploitation for the fur trade is the leopard cat, and then onlyin China. A CITES project is currently underway to
study the ecology of the leopard cat to aid evaluationof its status, and to examine harvest practices in China
in order to advise the government on development of asustainable management program (Johnson and Fuller
1992, Johnson et al. 1993; see Project 13 in Part III).
2. The North American example has demonstrated thatcommercial trade in cat furs can be sustainable, and canhelp to conserve cat habitat as part of a strategy of making wildlands an economically competitive form ofland use. Management strategies should continue toinclude funding for a strong component of research, andshould continue to evolve to reflect new understandingof species biology (Project 98).
3. Illegal trade in pelts persists in many range states, but
the areas which produce garments made from protected
species are few in number, and the customers appear
to be mainly tourists. With regard to the sale of cat fur
coats in Kathmandu, Van Gruisen and Sinclair (1992)
have pointed out that a small-scale publicity campaign
aimed at tourists would be a relatively simple to carryout, and could be highly effective.
4. The Oriental medicine trade poses a grave threat to thetiger, and commercial poaching could well lead to its
virtual extinction in the wild within a decade. Antipoaching protection for tigers should be increased, butmarket forces will probably continue to dictate the status of wild tiger populations. Project 12 includes trade
surveys to provide better understanding of market organization and consumer demand, so that effective strategies for bringing the tiger bone trade under control canbe developed.

243

Part II: Major

Issues.

Chapter

5. Cats in Captivity

Part II
Major Issues in Cat Conservation

Chapter 5
Cats in Captivity
cat’s jawbone amidst ruins in Cyprus dating from about
6,000 BC. No cats are native to the island, and the findindicates that the first human colonists brought their catswith them (Serpell 1992).
Domestication was probably a gradual process at first,
with wildcats developing a symbiotic relationship withhumans based on rats and mice, which then as now would
have been pests in homes and stores of food (Zeuner 1963,
Serpell 1988). It would be difficult to say when thedomestic cat became fully tame, or adapted to life in captivity rather than in the wild. Perhaps it hasn’t yet: thehousecat has been termed the only “semi-domesticated”
mammal in recognition of its retained ability to feed itself,
as well as its notorious reluctance to obey commands(Clutton-Brock 1988). While the housecat’s behavior has
not been totally altered by a captive environment, selective
breeding has produced great changes in its appearance.
Most of this “un-natural selection” has taken place over thelast century (Serpell 1992): Darwin (1868) mentioned that
there were only two distinctive types of domestic cat in
Britain (the Persian and the Manx), but there are now 27
officially recognized “races” and 3,667 color varieties(Feline International Federation in Zitt. 1993). The morphological, behavioral, and genetic differences between
domestic cats and wildcats deserve further study, andcould help to illuminate the effects of captivity on otherfelid species kept for future reintroduction to the wild (seeChapter 6, Reintroduction).
Apart from the notable example of the domestic cat, thehistorical record is sketchy with regard to small cats in captivity. The ancient Egyptians appear to have trained jungle
cats to hunt without really domesticating them (MorrisonScott 1952). Marco Polo (1299) reported that Kublai Khan
kept hunting caracals, as did some Indian princes, a practice which continued in India into the early 20th century(Allen 1919a, Sharma and Sankhala 1984).
Big cats have long been popular in captive collections.
The Romans displayed lions, tigers, and cheetahs inmenageries, pageants, and arena combat (Toynbee 1973).
Cheetahs were tamed and trained to hunt by early rulers

Introduction
Cats have been kept in captivity for thousands of years,
and have served various human needs, some utilitarian(such as hunting rodents for pest control, or antelope for
sport) and others less so (the use of cats as pets, emblemsof status, or as a spectacle for public entertainment). They
have always been among the most popular exhibits inzoos. It is only recently, however, that keeping cats in captivity has come to be perceived as something which not
only benefits humans, but also cats themselves. Interestin contributions which the ex situ (off-site-i.e.,
in captivity) zoo community can make to in situ (in the wild) conservation has greatly expanded over the past two decades.
Major zoos are evolving from menageries, whose animals
were collected chiefly for entertainment and display, toconservation centers, with an important role to play in public education and wildlife conservation (Fig. 1). In termsof conservation, zoos generally place the greatest emphasis on their efforts to maintain viable populations of threatened species, which can potentially serve as seed stock
for reintroduction in case of extinction in the wild, or for
revitalizing depleted populations.
This chapter reviews the status and management of cats
in captivity with specific reference to conservation of catsin the wild. Ways are recommended for the zoo community to strengthen its role in cat species conservation.

A Brief History ofCats in Captivity
The most obvious testament to cats’ historical popularity
with humans is the housecat, which was domesticatedfrom the African wildcat. Ancient Egyptian art reveals
that cats were fully domesticated by 1,500 BC, but thedomestication process clearly got underway much earlier,although there is little archaeological evidence (Pocock
1907c, Clutton-Brock
1981, Davis 1987, Serpell 1988,
Hemmer 1990). Recently, archaeologists have found a

244

Part II: Major

Issues.

Chapter

5. Cats in Captivity

Evolution of Zoos
ConservationCentre

Environmental
Resource Centre
Theme:

Environmental

Concerns:

Survival of species
Holistic conservation
Organizational networks I
Immersion exhibits

Exhibitq:
&&&d

Park

,
, a\_ ‘,

:,,,,
?

“”

“““’

’

LivingMuseum-”Theme:
Subjects:

EcOlOgiCal
Habitats of animak
Behavioral biologyS
ies managementIevelopment

8”

Exhibitty:>, -Concerns:

Species husbandry
Species propagation

Exhibitry:cages

DOramaS0

Figure1. Evolutionof Zoos.

Adaptedfrom G. Rabb’s

figurein WRl/lUCN/UNEP

of Palestine, Egypt, Assyria, Persia, Russia and theCaucasus(Vereschagin 1959,Guggisberg1975),and also,
according to Marco Polo, by the court of Kublai Khan.
This tradition reached its zenith during the time of the
Moghul emperorsin India ( 1500-1800s)(Divyabhanusinh
1984). One 16th century emperor,Akbar, wasreported byhis son, Jehangir, to have kept 9,000 cheetahsin his lifetime (Abu Faz-1 17thcentury), with over 1,000held at one
time in his menagerie (Alvi and Rahman 1968). Tigers
were a popularanimalin aristocraticcollectionsin Asia for
centuries(Courtney 1980),and may alsohave beentrained
to hunt by early Chineseemperors,long before the Mongol
invasion (Guggisberg 1975). Big cats have at times been
obliged by the powerful to serve as executioners. The
practice was not limited to the Romans-tigers were so
employed by Asian rulers, as well as pumasby the Incas(Guggisberg1975).
While peoplewere proficient at keeping cats in captivity, breeding them was a more difficult task. Regarding
both Akbar’s and his own collection of cheetahs,Jahangir
noted, “He was very eagerthat they should pair, but thisin no way came off. Several times the male and female

1992

ChicagoZoologicalSociety(1992).

hunting leopardswere brought together unchainedin thegardens. But there too they did not pair. Recently it happened[after Akbar’s death] that a male slippedoff its collar, approacheda femaleand paired with it. After two anda half monthsthree cubswere born. They grew up. As it
appearedstrangethe event wasrecorded.” He madea similar observation regardingAkbar’s collection of tigers: “Itwasjust a freak that a tigress becamepregnant and after
three monthsbore three cubs. Else, it hasnever happened
that a wild tigresspaired with its male in captivity” (Alviand Rahman1968,D. Chavda in Zitt. 1993).
Lack of successfulbreedingin captivity wasprobably amajor causeof the extinction of the Indian cheetahby theearly 20th century, as live-trapping of cheetahs was amajor industry in the late 1800s (Sterndale 1884,
Divyabhanusinh 1984). An additional factor wasthat only
wild-caught adult cheetahshunted successfully, but not
juveniles. The Alipore Zoo first imported an African cheetah in 1892, princely imports began in 1918 (D. Chavda
in Zitt. 1993), and by the late 192Os,Finn (1929) reportsthat import of African cheetahsfor the royal collections
had becomea regular practice.

245

Pat? II: Major

Issues.

Chapter

5. Cats in Captivity

Whereaszoos and menageriesthroughout history wereprimarily stockedby wild-caught animals,the entry into
force of the Convention on International Trade inEndangeredSpeciesof Wild Flora and Fauna (CITES) in1975(seeChapter4) meantthat from then on international
trade in most speciesof wild cat was prohibited, except
under very special circumstances. Public exhibition isconsidereda commercialuse, and so zoos beganto focus
on ensuringthe sustainability of their captive populations
through breeding programs, which involve inter-zoo
exchangesof individuals selectedaccordingto their bloodline. Over 90% of new animals registered in ISIS zoos
(see below) are now obtained through captive breeding(Flesnessand Foose 1990).

.
Table 1World Zoo Associations
Africa (15 zous)
Regional association: Pan African Associationof Zoological Gardens, Aquariums, and Botanic
Gardens (PAAZAB)

Asia (545 ZOOS)
National associations in: China, India, Indonesia,
Japan, Pakistan, Thailand
Regional association for southeast Asia: South
East Asian Zoo Association (SEAZA)

Status of Captive Populations

Australasia (30 zoos)
Regional association for Australia and New
Zealand: Australasian Regional Association of
Zoological Parks and Aquariums (ARAZPA)

There are probably well over 10,000 zoos around theworld (institutions which exhibit non-domesticanimalsto
the public: IUDZGKBSG 1993). At present, however,comprehensivedataare availableonly from approximately
1,100 zoos organized into national, regional, or international federations.Thesefederationsandtheir membershipare shownin Table 1.
Data on catsheld in zoo collections(Fig. 2) are derived
from the International SpeciesInformation System (ISIS),
which contains data on specieskept by over 400 zoos
(December 1992). ISIS data are incomplete-less than
40% of the organized zoos register their collections, andthere are problemswith irregular reporting. ISIS data arebiasedtoward large zoosin developedcountries,anddo notinclude information from specializedprivate breeders.For
example,in comparisonto the numberspresentedin Figure
2, the 1991 InternationalCheetahStudbook (Marker-Kraus
1992)reports a captive population of 1,006, and the Tiger
Global Animal Survival Plan (Tilson et a2. 1992) reports
1,075tigers. Similarly, P. Quillen (in litt. 1993)reportsthat
thereis a significantpopulation of oncillasin private hands
and in non-ISIS zoos. However, only ISIS data have beenusedin order to be consistentregarding the relative representation of species in captivity.
The World Zoo
Conservation Strategy (IUDZGKBSG 1993), which recommendsways in which the zoo community canexpandits
role in speciesconservation, emphasizesthe necessityof
improving data collection within the community.
Examination of the data shows that, with regard tospeciesvulnerability, the statusof captive populationsparallelsthe statusof researcheffort describedin Chapter3, inthat the big cats are disproportionately represented. The
average size of captive populations of the eight species
weighing over 20 kg is 491, ranging from a minimum of
144 (clouded leopards) up to 887 (tigers). The average
population size of the remaining21 speciesis 72, ranging

Central and South America (100 zoos)
National associations
Mexico, Venezuela

in: Brazil, Colombia,

Regional associations: Central American Zoo
Organizationand Latin American Association of
Zoological Gardens and Aquariums

Europe (300 zoos)
National associations in: Britain and Ireland,
Sweden, Denmark, Netherlands, Germany(plus Austria and Switzerland),
Poland, Italy,
France, Spain, Hungary
Regional association for Europe: European
Association of Zoos ancj Aquariums (EAZA)North America (175 zoos)
National associations

in: Canada

Regional association for the subcontinent:
Zoo and Aquarium Association (AZA)

AmericanTotal: 1,I 65 zoos

At the global level there is one single zooorganization:IUZDG-theWorld Zoo Organization.
Sources:

CBSG 1993, IUDZGKBSG(1993)

246

Part II: Major

Numberof SmallCats (<20 kg) held in Zoos
II

250
v)
a
4
3
F

200
150

5
&

100

e
z”

50
0
serval

co
3
g
.-z
z
z
f

bobcatcaracalE. lynx

Numberof SmallCats(< 20 kg) held in Zoos

wildcatfishingcat

CanadaNumberof SmallocelotReportingleopardcat

to ISIS in 12192

100
75
50
25

z”
0
lynx

junglemargayCats (c 20 kg) held in Zoos

cat

Reportingjaguarundi

black-footed

cat

cat

goldencat

to ISIS in 12/92

100
24
2.>
5
E

50

ii
f

25

75

z’
0
spottedcat

cat

Numberof Large

goldencat

catCats (> 20 kg) held in Zoos

Reportingto ISIS in 12/92

lion

snow
leopard

puma

1000

;

800

5
5
5

600

$
E
z’

400

200

0
tiger

leopardcheetah247

jaguar

Chapter

5. Cats in Captivity

Figure 2. Representationof felid species
in zoocollectionsin December,
1992.
Source: Jill Mellen, Co-chair,
Felid Taxon Advisory
Group,
AmericanZoo and Aquarium
Association.

to ISIS in 12/92

ReportingIssues.

cloudedleopard

Part II: Major

Issues.

Chapter

5. Cats in Captivityof these 60% were big cats of the genus Panthera (including the snow leopard). A survey to determine the amountof space available to carnivore species in North American
zoos (Mellen et al. 1992) shows that cats occupy about
50% of carnivore cage space (n=l,889 spaces), and arealso projected to occupy a similarly disproportionate share(47%) of spaces to be added within 5-10 years (n=322
spaces). Within the cat family, more than 70% of current
and future space is allocated to cats weighing over 20 kg
and a substantial portion is taken up by relatively common species (e.g., lynx, bobcat, puma).
Like wild habitat, captive habitat, or enclosure space,is in limited supply. Various estimates have been put forward as to how many viable populations of species could
be supported in the world’s captive habitat-which
altogether amounts to only some 80 km2 (Wiese and Hutchins1993). These estimates generally indicate that there isroom for no more than 1,000 species in viable numbers ofat least several hundred individuals (Soule et al. 1986,
Flesness and Foose 1990, Wiese and Hutchins 1993).
However, again the cats are allotted more than their “fair”
share. Seal and Foose (1992) estimate that the felid sections of the world’s organized zoos have the capacity tohouse at least 11,000 cats, and possibly double that. If thelower figure is accepted, this would permit maintenance of
36 species in populations of 300 or, looked at another way,
220 subspecies in populations of 50.

from 210 servals to two African golden cats. The vulnerable small to medium-sized cats (Categories l-3) are
under-represented.
Six out of 13 species are not kept in
captivity according to ISIS data (Table 2), and the average population size of the seven that are is just 30. With
the exceptions of the fishing cat and black-footed cat, there
are really no significant ex situ populations of these cats.
Study of these species is a priority recommendation ofthis Action Plan. Knowledge of the ecological requirements and status of these species will help to determine
whether any individuals should be removed from the wild
for the purpose of establishing “insurance” populations in
captivity.
International trade in live cats also parallels their representation in captivity (Table 3). That annual trade in
the big cats is so high, particularly for the tiger and lion,
reflects the size of the “unofficial”
captive populations
kept outside the major zoos and the breeding success thathas now been achieved. Much of the trade in these two
species involves travelling circuses and menageries(WCMC unpubl. data).
Another problem facing the management of ex situ populations is that most captive breeding programs follow
guidelines which segregate animals according to established geographic origin or to recognized subspecies.
Individuals of questionable status-“genericanimals”should technically receive lower priority for breeding than
individuals of known origin (Ryder et al. 1988, Wildt et al.
1992a), although Willis ( 1993) points out that this could
lead to a decrease in genetic diversity. The issue of recognition and classification of intraspecific diversity continues
to be debated within the zoo community, and is discussedwith regard to wild populations in Chapter 3. Table 4shows that a high percentage of individuals from most captive cat populations are generic animals. Out of 28 species
maintained in captivity for which there are data on founder
origin, more than two-thirds (n=20) have a generic component greater than 60%. This problem is particularly
acute for the small cats, although the high figure for thelion is striking.
On the positive side, the well-represented bigger cats
are increasingly better managed; as discussed below, captive breeding and population management programs areat various stages of development for all these species.
Management of these species as global populations is very
important, for at present regional captive populations arevery diffuse, with individuals scattered among an averageof 126 zoos. Several of the smaller species are also coming under the management of these programs.
Moreover, for years members of the cat family havebeen fortunate enough to occupy the “lion’s share” of thelimited enclosure space available to Carnivores. Foose
and Seal (1986) showed that at the end of 198 1, about halfof the 5,640 carnivores registered in ISIS were felids, andReproduction
Speciesin Captivity

Bred in Captivity

Most cat species have been bred in captivity, but with varying levels of success. This information is presented inTable 5. Exceptions include the Bornean bay cat andAndean mountain cat, which have not-as
far as is
known-beenmaintained in captivity. The Chinese mountain cat is kept in several Chinese zoos, but has not bred

Table 2Cat Species Not
Represented in ISIS Zoos
Bornean bay cat, C. badia
Chinese mountain cat, F. bieiti
lberian lynx, L. pardinus
Kodkod, 0. guignna
Andean mountain cat, 0. jacobitus
Flat-headed cat, P. pianiceps

248

Part II: Major

Issues.

Chapter

5. Cats in Captivity-(Tan Bangjie in Zitt. 1993). The first breeding center for the
Iberian lynx has been established at the Dofiana National
Park in southern Spain. The founding population consistsof two young females (three years old) and an older male
(7-8 years old). These animals cannot be returned to thewild-thefemales due to injuries, and the male because
he was captured as a cub and raised in captivity. No breeding has taken place so far: although the females showed
obvious signs of estrous behavior in the spring, the male
showed no interest (M. Aymerich in Z&t. 1993).
The large cats, particularly lions and tigers, have generally reproduced well in captivity. For more than 15
years, many zoos have used contraceptives to limit reproduction of their big cats. The most common of thesedevices are synthetic progestagen (melangestrol and megestrol) implants. However, research (Munson and Mason
199 1, L. Munson in prep.) is linking these implants to uterine lesions that may cause permanent infertility (severe
endometrial hyperplasia and endometrial polyps) or evendeath (uterine cancer).
Reproduction in captivity has been problematic for twoof the bigger cats: the cheetah and the clouded leopard.
Developing guidelines to improve reproductive performance in cheetahs has been the subject of a good deal of
focused research as well as wide-ranging surveys over the
past few years (Grisham 1988, Marker-Kraus et al. 1990,
Lee 1992, McKeown 1992, Lindburg et al. 1993, FitchSnyder in press, N. Wielebnowski
in prep., S. Wells in
prep., J. Grisham in prep.), including a major effort by the
Cheetah Species Survival Plan to assess the reproductive
status of 128 captive cheetahs in North America (Wildt etal. 1993a). This effort is bearing fruit: the number of successfully reproducing cheetahs in North American zoos
has increased by 66% (to 53) since 1986. Of all cheetah
births since 1956 in North America, more than one-third
have taken place since 1986 (Marker and O’Brien 1989,
Marker-Kraus and Grisham 1993).
Of the 79 clouded leopards listed in the International
Clouded Leopard Studbook in 1986, less than 20% of adultfemales and 27% of males had reproduced (Wildt et al.
1986a). Research into the management and husbandryrequirements of the clouded leopard is only just beginning, but the main problem is mate incompatibility, often
fatal. The best solution seems to be establishing pairs when
the animals are still sexually immature (Yamada andDurrant 1989).
In general, reproduction of the smaller cats (<20 kg) in
captivity has been inconsistent (Mellen 1989, 199 1, 1992,
1993). Many regional captive populations are highly
inbred (Mellen 1989, 1993; G. Foreman, P. Quillen, pers.
cornm.), including Geoffroy’s cat, rusty-spotted cat, pampas cat, fishing cat, wildcat, and African golden cat. In a
study examining environmental components as potential
correlates to reproductive success, 20 species of small cats

Table 3
CITES-reported International
Trade in Live Cats, 19764990

Trade
Tiger, R tigp&
tigrisLion, P. ka?
Leopard,R pardus
Cheetah, A. jubatus
Puma, I? concolor
Eurasianlynx, L. lynx
Jaguar, I? 0incd
Canadalynx, 1. canadensis~
camdensisf
Serval, L. sewal
Caracaf, C. caracal
Wildcat, F: sr’/ves&&
sikestrls
Snow leopard, CI.uncjd
Leopardcat, I? bengalens&
Bobcat, 1. rufus
Ocelot, 1. pardalis
Cloudedleopard, IV, nebulosa
Jungle cat, F. chaus
Black-footedcat, F. nigripes
Geoffroy’s cat, 0. geoffroyi
Asiatic golden cat, C. temminckr^
Manut, 0. manul
Sand cat, F. margarita2
Oncilla, L. tigrlnus
Margay, L. wiedi
Jaguarundi,H. yaguarundi
Fishingcat, P. viverrinus
Rusty-spottedcat, P. rubiginosus
Marbled cat, P. marmorata
Flat-headedcat, P. plar?iceps
African goldencat, I? aurata
Chinesemountaincat, F. bieti
Pampascat, 0. calocola
Kodkod, 0. guigna

239.8
145.1
82.8
61.U
26.5
23.9
23.1
22.8
20.7
20.4
16.3
15.5
13.1
10.3
8.2
8.0
5.0
3.8
3.7
2.9
2.4
1.9
1.9
1.7

1.5
1.5
0.8
0.7

0.5
0.5
0.2
0.2
0.2

1 The annual average is inflated because80 lynx
in I989 were exported from Canada to the U.S.
for reintroductioninto New York state. Canada
reports exporting 140 animalsto the U.S. in
1990;there may be an error in the data.
2 The annualaverage is inflated by a shipmentof
20 animalsfrom Pakistanto Germany in 1977,
Trade uver the remainingperiod amountsto
only eight animals.

249

Pat? II: Major

Issues.

Chapter

5. Cats in Captivity.
Table 4
Animals of Unknown

Subspecies/Origin

Species

Fishing cat, I? viverrinus
Pampas cat, 0. colocolo
Marbled cat, P. marmorata
Ocelot, L. pardalis
Jaguarundi, H. yaguarondi
Geoffroy’s cat, 0. geoffroyi
Jaguar, P. onca
Manui, 0. manul
Serval, L. serval
Caracal, C. caracal
Clouded leopard, IV. nebulosa
Lion, P. lea
Leopard cat, P. bengalensis
Jungle cat, F. chaus
Black-footed cat, F. nigripes
Oncilfa, L. tigrinus
Asiatic golden cat, C. temmincki
Bobcat, L. rufus
Puma, P. concolor
Margay, L. wiedi
Cheetah, A. jubatus
Leopard, P. pardus
Rusty-spotted cat, P. rubiginosus
Eurasian lynx, L. lynx
Tiger, P. tigris
Wildcat, F. silvestris
Canada lynx, L. canadensis
Sand cat, F. margarita

in ISIS Zoos in 1989

Vulnerability
Ranking

Total
Population

2
5a
3
5a
5c
4

25
IO
4
97
44
36
208
9
161
104
116
522
36
27
38
4
27
170
240
65
318
577
12
47
847
28
60
13

3(A)
4
4
5b

3(A)
3(A)
5b
5b
2
3
3
5a

5wv
4

3(A)
WA)

3
5a

2(A)
5c
4
4

housed in eight zoological institutions were evaluated(Mellen 1991). Variablesthat correlatedsignificantly withsuccessfulreproduction included housing no more than
one male and one female together; relatively few medical
problems (i.e., healthy cats); and a high level of keeperanimal interaction.
In a related study, the effects of hand-rearingon subsequent ability to copulate was examined (Mellen 1992).
Resultssuggestthat catswhich have beenhuman-rearedina traditional zoo nursery setting are lesslikely to copulate
successfullyasadults. On the other hand, adult reproductive successdoes not appear to be compromised in cat
speciesthat were human-rearedin a rich and varied environment (J. Mellen, pers. obs.). This suggeststhat thehand-rearing proceduresof zoos should be re-evaluated,and should include provision of companionsand a more

Pwwnt {Number]
Unknown
SpecieslOrigin
1ooo/o
100%
100%
96%
93%
92%
91%
89%
87%
86%
85%
79%
78%
78%
76%
75%
74%
73%
65%
64%
42%
29%
25%
17%
12%
7%
0%
0%

I251

WI
ELtl

P31
WI

WI
VW
PI
Et401
WI

WI
t4-141

E281
PII
E2Ql
I 31
f 201
VW

II561
WI

[I351
f 1691

PI
PI
[IO41
PI
-

“interesting” environment.
Zoos have madesignificant stridestoward establishingand maintaining self-sustainingcaptive cat populations.
However, zoosneedto increasethe representationof mostsmallcat species,and especiallythe rarer ones. The future
viability of presentpopulationsis tenuouswithout the addition of new animals, and it would be unwise to seekto
remove animalsfrom the wild before researchinto theirstatus and ecology has been carried out. Instead, there
should be more cooperation between zoos in different
regionsof the world in order to take full advantageof all
animals presently held in captivity. The program now
being established for tigers (see below: Tilson et al.
1992)-the culmination of many yearsof interest,data collection, and international communication-is a good
exampleof how this cooperationshouldunfold. Other ini-

250

Part II: Major

Issues.

Chapter

5. Cats in Captivity.
Table 5
Captive Pr topagation Status of Cat Speciesl
Specieslsred in
captivity

RaisedYoung

Zd Generatbn
BirthsSeffSUtstaZning32

Cheetah, A, jubatus
Caracai,c. cz?racal
Asiatic got&n cat, C. temmincki
Jungle cat, f. chaus
Sand cat, E margarita
Black-footed cat, E nigripes
Wildcat, F. sikestris
Jaguarundi, H. yaguarundi
Ocelot, L. pardalis
Oncitia, L. tigrinus
Margay, L. wiedi
Send, L. serval
Canada lynx, 1. canadensis
Eurasian lynx, L. lynx
iberian tynx, L. pardinus
Bobcat, L. rufus
Pampas cat, 0. ~olo~ok~
Geoffroy’s cat, 0. geoffroyi
Kodkod, 0. guigna
Manul, 0. manul
Leopard cat, P. bengalensis
Flat-headed cat, P. planiceps
Rusty-spotted cat, P. rubignosus
Fishing cat, P. viverrinus
African golden cat, P. aurata
Puma, P. concolor
Clouded leopard, N. nebulosa
Lion, P. lea
Jaguar, P. onca
Leopard, P. pardus
Tiger, P, tigris
Marbled cat, P. marmurata
Snow leopard, U. uncia

Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes

Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
Yes
Yes
Yes
Yes
Yes
Yes
No
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
Yes

Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
Yes
Yes
Yes

Yes
Yes
No
IMavbe
No*
No
Maybe
No
Yes
No
Maybe
Yes
Yes
Yes
No
Yes
No

Yes
Yes

NO

NO

NO

NO

Yes
Yes
No
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes

Yes
NO
No
Maybe
NU
Maybe
Maybe
Yes
Yes
Yes
Yes
NO
Yes

References

=f,6
133
l-t
If
12
-w
1,214
1,637
5
W
1
13
2
k2
139
I
1
1
1
1
1
1, IO
1

1 includesonly institutionsreportingto ISIS and/or monitoredby the internationalZoo Yearbook, with the exceptionof the newly establishedIberianlynx breedingcenter, Dofiana NationaiPark, Spain.
2 Sustainabilityassessedat specieslevel according to total populationsize (* 50) and our best informationor
guesstimateon numberof originalfounders.
3Generics only, no breedingprogram (A. Shoemakerin litt. 1993).
References
I. Seager and Demorest1978; 2. Mefien 1993; 3. Sausman1989; 4. Olney and Ellis 1991; 5. P. Quillen, pers.
comm.; 6. ISIS MammalAbstract 1991; 7. Anderson 1977; 8. Schaffer and Rosenthal1984; 9. Tonkin and Kohler
1978; 10. Barnes 1976; 11. J. Meilen, pers. comm.; 12. M. Aymerich, pers. comm.; 13. Olbricht and Schtirer 1993.
Source: Jill Meiien, Co-chair, Fetid Taxon Advisory Group, American Zoo and AquariumAssociation

251

Part II: Major

Issues.

Chapter

5. Cats in Captivity.
hble 6Structure of an AZA Species Survival Plan (SSP) &laster Plan
1. Jntruduction
4. introductory comments
3. Outline of objectives
1. Overall goal (e.g., retain 90% of genetic diversity for 200 years)
2. Number of founders desired, available, and presently in population
3. Carrying capacity
4. Generation time management
5. Management strategy (e.g*, by mean kinship)
6. Lifetime offspring goats (number of offspring needed per individual founder and non-founder)
7. Population control methods
8. II situ recommendations
9. Othersa. Special medical considerations
b. identification system for SSP animalsc. Policy for breeding recommendations
d. Special research goals
e. Surplus animal policy
Genetic Analysis
A. introduction
I. introductory comments (i.e., special considerations)
2. List of parameters and assumptions used in analysis
3. Overview of results of analysis
B. List of mean kinship coefficients
C. List of inbreeding coefficients
D. Gene drop output (i.e., model of rate of genetic drift)
E. N, estimations
Demographic Analysis
A. introduction
1. Introductory comments (i.e., special considerations)
2. List of parameters and assumptions used in analysis
3. Population summarya. Population numbers
b. RecentHistorical growth or declinec. Generation timed. Age of first reproduction and senescence
6. Life table (includes lambda, T, r, etc.)
C. Age pyramid
Recommendations
A. Institution by institution (sorted by institution)
1. Breedings for next few years
2. Transfers for next few years
3. Surplus
4. Animals to be held in non-breeding situation
B. Animal by animal (sorted by studbook number)
1. Breedings for next few years
2. Transfers for next few years
3. Surplus
4. Animals to be held in non-breeding situation
C. Other (e.g., research and medical)
AppendicesA. Tables
B. Graphs and figures
C. Other information

252

Pat? II: Major

Issues.

Chapter

5. Cats in Captivity

Table 7
Captive Breeding Programs and Studbooks few Felid Species
North America: Species Survival Plan (SSP)
Cheetah, A. jubatus (1984)
Clouded leopard, IV. nebulusa(I 989)
Asiatic lion, P. lea per&a (t 981)
Amur (P.t a&aica)+ Sumatran (P. f. sumatrae)tigers (1992); Indochinese
under development; Generic tigers
Snow leopard, U. uncia (I 983)

tiger (I?& corbe#i)

Europe and Russia: European Endangered Species Programme (EEP)
Cheetah, A. jubatus
Clouded leopard, IV. nebulosa
Amur (P.p. opienWs) + Persian (p.P, saxiculo~ leopards
Amur (I? t. figris) + Sumatran (P.t. Sumatra@tigers
Snow leopard, U. uncia

British tsles: Joint Management of Species Group (JMSG)
Cheetah, A. jubatus
Margay, L. wiedi
Ocelot, L. pardalis
Geoffroy’s cat, 0. geoffroyi
Clouded leopard, IV. nebulosa
Asiatic lion, P, lea persica
Persian leopard, (p.P. saxicohz) (1988)
Amur (Et. altaica) and Sumatran (I? f. sumatrae)tigers
Snow leopard, U. uncia

Indonesia: Indonesian Species Programme (PKSBI)
Sumatrantiger (P.t tigris)Australasia: Australasian Species Management Programme (ASMP)
Asiatic golden cat, C. femmincki(I 993)
Persian Jeopard (p.P. sz&u/@-may
be dropped
Sumatran tiger (I? L sumatrae)(I 992)

in favor of Javan leopard(P.p. me/as)(1993)

Japan: Species Survival Committee of Japan (SSCJ)
Cheetah, A. jubatus
Amur tiger, P. tigrisaltaica
Snow leopard, U. uncia

Studbooks (I=lnternational; R=Regional)
Cheetah, A. jubatus: l-1 982; R (Great Britain)-1 992
Sand cat, F. margarita:I-t 988, but unofficial since late 1970s
Black-footed cat, F. nigripes:t-1 988; R (North America)-under
Ocelot, L. pardalis: R (North America)-underdevelopment
Margay, L. wiedi: R (Great Britain)
Geoffroy’s cat, 0. geoH?oyi:R (Great Britain)

development

Continued

253

on next page

Pat? II: Major

Issues.

Chapter

5. Cats in Captivity

Rusty-spotted cat, P. ‘rubiginosus: l-under development
Fishing cat, P, viverrinus: R (North America) and l-f 993
Clouded leopard, IV. nekdosar t-1980; R (Europe, India)
African lion, I? lea: 1-l 992
species account);
Asiatic tiony~P. lea per&x I-1982 (suspended-see
development; unuff iciaf studbook pubi. 1988
Jaguar, P. onca: R (North America)-under
Asian leopard subspecies: North Chinese (P.p. japonensis), Sri Lankan (F?p, kotiya),
Amur (I?p. ot;ientaiis)
and Persian (P,p. saxjcoloi): I-1 977; R (Europe, Great Britain, Australasia: P.p. saxico~o~Amur tiger, Et. alfaka: f-l 976
Sumatran tiger, Et. sumatrae: I- 1978; R (N. America)-1990;
R (Indonesia) under dev&pment
Bengal tiger, P.t tigris: t-l 979; R (India)-under development
0 ,..rL 1 blltlla
llL:,1. ttyerr:,,, $ P.n 1.A awoytwsK5.
lu~l,--A:~, I-I 4f 4360
auutr
vo3
Indochinese tiger, P. t. corbetti; 1-I 990
Snow leopard, U. uncia: I-1 977

ioral traits did not become a major issue until the second
conference, held in July 1976 (Tudge 1976, Leyhausen1977), and interest has grown exponentially since then.
Captive breeding and population management programs became institutionalized when the American Zoo
and Aquarium Association (AZA) launched its Species
Survival Plan (SSP) program in 1981. Under this program, zoos cooperate to manage individual animals withina number of zoo populations as a single population. The
master plan includes demographic and genetic analysesof the population, based on studbook data, and makes recommendations for each animal in the population, includingwhich ones should breed and with whom; which ones
should not breed; which ones should be removed from the
population; and which ones should be used for research,
reintroduction, or genome banking (R. Wiese in Z&t. 1993).
The outline of a SSP master plan is presented in Table 6.
Such programs allow for the ultimate in hands-on management, population manipulation to a degree that would
be impossible to duplicate in the wild. Other regions have
now established similar programs, and the species being somanaged are shown in Table 7.

tiatives include technical assistance from experts in developed countries to zoos in developing countries in enclosure
design and husbandry, animal selection (of known origin,previously held by other zoos or by private owners) forbreeding programs, veterinary care and genetic analysis. In
exchange for their assistance, western zoos would acquire
access to new founders for breeding programs, and newmaterial for genetic research (Quillen 1992; W. Johnson, S.
O’Brien, P. Quillen, W. Swanson, pers. comm. 1993).

Captive Breeding and Population
Management
Programs
The first stage in the development of inter-institutional
breeding programs began in 1966, when international studbooks were organized under the aegis of the International
Union of Directors of Zoological Gardens (IUDZG) totrace the history of all individuals of rare species or subspecies in captivity (Dathe 1980). These registers areindispensable for tracking the degree of relatedness
between animals. Studbooks for selected tiger subspecies(Seifert and Muller 1976), snow leopards (Blomqvist
1978), and selected leopard subspecies (Shoemaker 198 1)
were the first established within the cat family. However,the studbook system essentially left it up to individual zoos
to take the initiative to act upon the demographic information contained in the registers and establish breeding
consortia for the purposes of animal exchange and managed breeding.
The establishment of more structured breeding programs was urged at the first international conference on
Breeding Endangered Species in Captivity, held in the
U.K. in May 1972 (Smith 1972). At that time, the focus
was on building up captive populations to avoid the needto remove more individuals from the wild. Concerns about
preservation of genetic diversity and wild-adapted behav-

A Global Captive

ActionPlan for Felids

A Global Captive Action Plan (GCAP) for Felids (Wildt etal. 1992a) was produced in 1991-1992 during two consecutiveannualworkshopsheld by the AZA’s Felid Taxon
Advisory Group (TAG). The Felid TAG holds annual
workshopsto review progressand revise the Plan asnecessary. The GCAP prioritizes speciesby region for theestablishmentof viable captive populations according torarity in the wild. The Felid TAG workshops haveattemptedto quantify in situ rarity by applying the MaceLande criteria (Mace and Lande 1991) to 259 subspecies.
The GCAP’s priority ranking appearsin Table 8, with each

254

fart

II: Major

Issues.

Chapter

5. Cats in Captivity

Table 8A Global Captive Action Plan for Felids (Wildt et al, 1992a)l

Cat Action Plan
Regional VulnerabiMy Ranking*Gbbaf CaptCve Action Plan
Ranking (I = highest prkwity)
Africa1. Cheetah, A, jubatus
2. Black-footed cat, F. nigripes
3. Lion, P. lea
4. African golden cat, P, aurata
5. Serval, L. serval
6. African wildcat, F.s. lybica group
7. Caracal, C. caracal

N=l (A); S=2(A)I
2(A)
1
N=2(A); S=3
s=5

S=4; N=Sa (A)Asia
1. Tiger subspecies:altaka, amuyensis,corbeR&sumatrae
Iriomote cat, P. [b=Jiriumotensis
2. Asiatic lion, P. lea persica
3. Flat-headedcat, P. planiceps
Bornean bay cat, C. badia
Marbfed cat, P. marmorata
Snow leopard, U. uncia
Clouded leopard, IV. nebulosa
Bengaltiger, P.t. tigris
4. Manul, 0. man&
5. Asiatic golden cat, C. temmincki
6. Chinesemountaincat, F. bieti
7. Persianleopard, P.p. saxicolor
8. Amur leopard, P.p. orientalis
9. Sri tankan leopard, P.p. kotiya
IO. Asiatic wildcat, F.s. ornata group
1I. North Chineseleopard, P.p. japonensis
12. SouthvvestAsian wildcat subspecies
EuropeI. Iberian lynx, L. pardinus
2. Scottish wildcat, F.s. grampia
3. Europeanwildcat,,Fs silvestrisgroup
4. Eurasianlynx, L. lynx

w

no ranking, but high priority
1
2

1
2
1(A)
2(A)
l(A)
3
2

1
3(A)
3(A)
3(A)
As=2
3(A)
N=5b

1
Er=3
3
Er=2; As=3

North America
1. Florida panther, P.c. coryi
2. Texas ocelot, L.p. albescens
3. North American puma subspecies
Canada lynx, L. canadensis
Bobcat, I. rufus
4. Central American puma subspecies

4(A)

4

4(A)
3
4

4(A)

Latin America
1. Andean mountaincat, 0. jacubitus
2. Kodkod, 0. guigna
3. Oncilla, L. tigrinus
4. Margay, L. wiedi
5. Pampascat, 0. coloc0~0

1
1
2

3
4
Continued on next page255

Part II: Major

Issues.

Chapter

5. Cats in Captivity

Global Captive Action Ptan
Plan
Ranking (1 = highest priority)

Cat A&cm Plan
Regional Vulnerabiiisy Rankings

6,
6. Texas jaguarundi, Hy, cacomjflj
7. Ocelot, L. pardalis
pardais8. Jaguar, P. oncamm
9. Geoflroy’s cat, 0. geo&uyi
IO, Jaguarundi, H. yaguarondi

Nut included in the GCAP
T=4; N=fia
452
2

Jungle cat, F. chaus
Sand cat, F. margarita
Leopard cat, P. bengalens&Rusty-spotted oat, P. rubiginosus
Fishing cat, P, viverrinus

1 As explained under Taxonomy, this Action Plan generally avoids reference to subspecies, given thequestionable validity of traditional subspecific designations and the absence of modern taxonomic
investigation of subspeciation fur
for most cats. The regional vulnerability rankings used in this Action Pian
do not necessarily correspond to the subspecific classification used for the GCAP.
As=Asia sub-region
ErkEurope sub-region
N=North Africa and Southwest Asia
S=Sub-Saharan Africa
T=Tropical Asia

felids or carnivores (CBSG 1993). Tables 9 and 10 present
the North American and Australasian RCPs, which establish regional priorities not only according to the GCAP, butalso according to practical considerations, including captive space limitations within the region, number of knownorigin captive animals, need for taxonomic study at the
specific or sub-specific level, presence or absence offounder stock, and the existence of other captive breeding
programs outside the region. Therefore, they look quite
different from the GCAP list of priorities, and include recommendations for reducing populations of certain species
while others are expanded.
While the GCAP considers multiple taxa, a Global
Animal Survival Plan (GASP) is concerned with just one
taxon, typically a species. A GASP sets out a strategy tocoordinate the activities of regional captive breeding programs. Within the family Felidae, a GASP has been produced only for the tiger (Tilson et al. 1992). The goals of
the Tiger GASP are shown in Table 11, and it is clear thatthe objective is to integrate ex situ and in situ conservation efforts for the tiger to a greater extent than any felid
captive breeding program has attempted before.

species’ vulnerability ranking given for comparison.
Although global in scope, the priorities currently tend toreflect the North American membership of the Felid TAG,
in that North American and European species are given
relatively high profile, while Eurasian, tropical Asian andsouthwest Asian species are lumped together. However,the GCAP is expected to change as new information on thestatus of species in the wild becomes available-for example, upon publication of this Action Plan.
The Global Captive Action Plan is based solely on thepremise that captive breeding programs should be established first for those taxa which are rarest or most threatened in the wild (Seal et al. 1993). It is recognized as an
idealized strategy that does not take practical difficulties
into account, such as the fact that founder captive populations of most rare small cats are very low or non-existent.
The GCAP is meant to serve as a guideline for implementation plans, including Regional Collection Plans (RCPs)
and Global Animal Survival Plans (GASPS-discussed
below). The AZA created the TAG groups for the purposeof drawing up RCPs at the family level; British, Europeanand Australasian zoos have also recently created TAGS for

256

Part II: Major*
Table 9North American 1993-1994 Regional Collection

Issues.

Chapter

5. Cats in Captivity

Plan for Felids(J. Mellen and D.E, Wildt in liilt. 1993) (nut in order of priority)

Recommend no breeding; reduce totai population to a target levee of no more than 30 animals to be held for
educational and research purposes only; assess SPARKS* database for population paramete&I

L. serval
Recommend no breeding; reduce total population to a target level of no more than 30 animals to be held for
educational and research purposes only; recommend propagating animals of known subspecies onfy.
Develop an international studbook for Asian subspecies and a regional studbook for African subspecies; reduce
total population to a target level of 50 African animals to be held for educational and research purposes only;
establish an Asian population of 25 pairs to eventually replace African animals; recommend propagating animalsof known subspecies only.
I

Pmrubigjnosus
Acquire founders; establish and expand population to 25 pairs.
E n/gripes
Establish regional studbook and eventually an SSP; establish and expand population to 25 pairs.
L. ti..riiws
Encourage an international studbook; establish and expand population to 25 pairs.
0. manul
Acquire additional founders; develop husbandry and propagation protocols; expand population to 25 pairs.

P, viverrinus
International studbook in place; acquire founders; establish and expand population to 25 pairs.
F. margarB
international studbook in place; expand population to 25 pairs.
L. pardalis
Estabtish a regional studbook; recommend propagating animals of known subspecies only; acquire new founders;establish and expand population to 25 pairs; maintain and expand relationship with the Texas ocelot recovery program(Feline Research Center, Texas A&J Univ.) and support a captive breeding program for this subspecies (if necessary).
P. leoCombine Asiatic and African lions under one SSP; recommend breeding for those animals of known origin only.
Recommend no breeding pending studbook review; identify studbook keeper.
P. pardus
Recommend no breeding of generics; see International Studbook on Rare Leopards for potential breeding stock.
A. jubatus, N. nebhsa,
SSP in place.

P. tigris, U. uncia

C. temmlncki, F. chaus, F. silvestris, H. yaguarondi, L. Medi’, 0. colocolo, 0, geoffroyi, P. bengalensis,
i? planiceps, P. aurata, P. marmwata
Not currently recommended for breeding or maintenance in North American institutions.
1 Except Florida panther.
2 SPARKS is a software system developed by MS to synthesize data from ISIS studbook-like reports.
3 Population parameters include size of population, age structure, number of institutions holding these animals,
among others.

257Part II: Major

Issues.

Chapter

5. Cats in Captivity*
Table 10Australasian 1993-l 994 Regional
Collection Plan for Felids

Table 11
Goals of the Tiger Global
Animal Survival Plan

(Christie(Tilson et a/# 1992)

1993) (nut in order of priority)

4.. pardalis
Replace with high priority SE Asian small carnivore.

lC. femmincki
Continue SMP*, expand holdings.
A. jubatus, P, tigris sumatrae
Continue SMP.

0 To contribute to tiger conservation by:* Developing, maintaining, and using captive
breeding programs to provide a genetic
and demographic reserve to re-establish orrevitalize wild populations when the needand opportunity occurs;/K ne&u!usa, P. leu persica
Acquire and develop SMP.
P. pardus saxicobr
Under review, develop SMP

To develop captive programs fur tiger
conservation with the paramount purpose ofcontributing to the survival and recovery of all
distinct taxa in the wild.

.

lP. pardus melas (Java)
Under review.
P, lea, lJ. uncia
Maintain.
C. caracal schmitzil’ (Turkmenistan),
Maintain two pair.

P. viverrinuslC. caracal, L. serval, LS lynx, L. rufus,
P. bengatensis, P. concalor, P. pardus
Remove from active management and phaseout holdings.

lConducting problem-oriented research thatwill contribute to management of tigers inboth captivity and the wild; collaborating onsuch research where appropriate with fieldresearchers; communicating and transferring
the results of such research to managers ofother captive and wild populations;
Providing where possible financial as wellas technical support for in situ Adopt-A-Parkprograms;
Using the tiger as a focus for conservation
education, public relations, and marketing
programs.

*Species Management Plan

Advancesin Assisted

ral patterns in gamete transport, implantation, gestation,
parturition, postpartum fertility, and age at first and last
reproduction. Fortunately, there are excellent baseline
data available from years of study into the reproductive
biology of the domestic cat (Wildt 1991a). Success withassisted reproduction in the domestic cat (Goodrowe etal. 1988, Howard et al. 1992) also provides model data
which can potentially short-cut similar experimentation inother species (Wildt et al. 1986b). However, some reproductive parameters can differ quite sharply even between
closely related species (Wildt et al. 1988, Brown et al.
1993), but with captive breeding programs as the catalyst,
a substantial database is gradually being built up about
the reproductive biology of different species (Wildt 1992b,
Wildt et al. 1993a).
Of key importance is accurate information on theendocrine status of the female. Fresh or thawed sperm
must be deposited coincident with ovulation, and embryos
must be placed into the recipient at a time when the uterus

Reproduction

During the last decade, there has been growing interest in
applying assisted reproductive technology, largely developed for domestic livestock, to wild animals. In the broadest terms, the technology includes artificial insemination,in vitro fertilization, embryo transfer, and gamete/embryo
cryopreservation.
While natural breeding of compatible
pairs and maternal rearing of offspring are the preferred
method of captive reproduction, and more effort needs tobe directed toward appropriate husbandry and behavioral
enrichment to ensure that this takes place, artificial propagation is a useful tool that also has potential application
toward maintaining the viability of in situ populations.
To be successful, assisted reproductive techniques rely
upon detailed understanding of fundamental reproductive
events, a considerable list including the following: seasonal or genetic impacts on reproduction, duration of the
estrus cycle, timing of ovulation, ejaculate norms, tempo-

258Part II: Major

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5. Cats in Captivity

as IVF embryos, were born to a Siberian tiger dam after
embryo transfer (Donoghue et al. 1990). The Asiatic wildcat kittens were produced in a domestic cat female (Pope etal. 1989). While this has been described as interspecifictransfer, the wildcat is very closely related to the domestic cat and is considered conspecific here. The exciting
aspect of interspecies embryo transfer is that “rare”
females could be hormonally stimulated to produce many
eggs, which could be fertilized in vitro and transferred to“common” female surrogates (Wildt 1991~). However,there is growing evidence that biological compatibility
between the trophoblast (progenitor of the placenta) of
the embryo and the uterine endometrium is a very complex
and restrictive phenomenon, and interspecies embryotransfer is unlikely to become routine (Wildt 1991b).
Also, IVF using frozen-thawed embryos has only beensuccessful with domestic cats (Dresser et al. 1988).
Finally, interest is growing in the establishment ofgenetic resource banks to hold frozen gametes, embryos,
blood products, tissue, and DNA collected from a large
number of species. Wildt (1992a) has envisioned genetic
resource banks as functioning much like a large financial
institution, with constant “withdrawals”
and “deposits”
performed by many branches and between continents. As
assisted reproductive technology advances, the advantagesof moving genes via germ plasm rather than living animals
are obvious in terms of time, money, simplicity, andsafety. Improved access to biological samples taken fromwild animals will also help to increase fundamental knowledge of species biology by providing more opportunity
for the participation of laboratory scientists in conservation
research (Wildt 1992a).
The tiger is the subject of the first Genome Resource
Banking Action Plan (Wildt et al. 1993b). The plan,
drawn up for the North American region, identifies key
captive animals for sperm, tissue, and blood collection, andrecommends collection of germ plasm from wild animals
in cooperation with field scientists. It represents the firstorganized effort to sample, evaluate, cryopreserve, catalog,
maintain and use wildlife germ plasm. The managers statethat the “resource of frozen tiger semen (and eventually
embryos) will be used interactively with living populations
to periodically infuse genetic material among diverse captive or wild stocks or preserved genes from previous generations” (Wildt et al. 1993b: 1).

is synchronous with the stage of embryo development. An
ability to trace endocrine patterns also provides the opportunity to diagnose pregnancy and predict parturition. The
standard method of endocrine monitoring involves regularly measuring hormone levels in blood, which may bepractical for captive animals, but not for wild animals:
anesthesia or immobilization stress can affect hormone
levels. Recent attempts to detect hormonal metabolites
non-invasively in animal wastes, both urine and feces,offer hope for development of a monitoring regime suitable to field conditions (Wildt 199 1b, Brown 1992).
However, the use of assisted reproductive technology
for wild populations lies well in the future. The techniques
will first have to be perfected in captivity, and their regularapplication will be most immediately useful to improving
the efficiency and effectiveness of captive breeding programs. These programs have run into problems when two
animals recommended for breeding turn out to be incompatible. In such cases, it would be simpler to use assisted
reproduction than to attempt to manipulate an array ofenvironmental factors to stimulate natural copulation. It
will also be less dangerous, more cost-effective and morepractical to transport frozen spermatozoa and/or embryos
between zoological institutions than stress-susceptible live
animals (Wildt 1992a, Ballou 1992, IUDZGKBSG
1993).
In relation to other taxa, there has been a relatively large
amount of research aimed specifically at refining assisted
reproduction techniques for captive felids. Artificial
insemination has been used to produce living young in theleopard cat, cheetah, tiger, puma and clouded leopard(Howard et al. 1992a, Wildt 1992b. Donaghue et al. 1993).
A litter of leopard cats has been produced using sperm collected and frozen at one institution and inseminated into afemale at another institution (J. Howard and D. Wildt
unpubl. data). All of these successes have relied upon theuse of exogenous hormones to stimulate ovarian activity,
followed by surgical laparoscopic deposition of sperm
directly into the uterine lumen. Vaginal or intracervicalinsemination in felids (other than the domestic cat) has
worked only once: a leopard cub was produced by thismethod and carried to full term, although it was stillborn(Dresser et al. 1982). The problem appears to be related toanesthesia, which relaxes the uterus and reduces the contractions which normally assist in sperm transport (Wildt
1991c, Howard et al. 1992b).
Through in vitro fertilization (IVF), embryos have been
produced for several felid species, including the domestic
cat, Asiatic wildcat, jungle cat, African golden cat, fishing cat, tiger, puma, snow leopard and cheetah (Goodrowe
et al. 1988, Donaghue et al. 1990, Miller et al. 1990, Popeand Dresser 199 1, Pope et al. 1989, 1993). The transferof these embryos into surrogate females has resulted in living young in the domestic cat (Goodrowe et al. 1988),
Asiatic wildcat, and tiger. Bengal tiger cubs. conceived

Linking Ex situ and In situ
Cat Conservation
There is a strong movement in the zoo community toward
becoming more closely involved with wildlife conservation (Anon. 1974, IUDZGKBSG
1993). The contribution

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Chapter

5. Cats in Captivity

The problems of reintroducing cats into the wild are
discussed in Chapter 6, but can be summarized here bynoting that, in general, it is not high on the list of priorities for cat conservation in the near future. For reintroduction, efforts should first be directed toward alleviating
the pressures which drove the population to extinction in
the first place. Otherwise, reintroduction is unlikely to be
successful, and may harm the cause of species conservation rather than advance it.
In terms of restocking, or infusing genetic diversity,
there are several major tasks which need to be accomplished before such action can be undertaken. First, small,isolated and endangered populations need to be identified
systematically through greatly increased field survey
efforts. Second, their demography and dynamics need to
be understood so that appropriate candidate populationsand possibly even candidate individuals-canbe selected.
Third, it is an open question whether new, captive-rearedanimals can be introduced to existing wild populations andboth survive and breed. Fourth, assisted reproductive techniques need to be further developed before they can beused effectively in the field.
The zoo community often predicts that wild populations will some day be as intensively managed as zoo populations (e.g., Tilson 1992b), but this will happen only if
it is accepted as inevitable that cat populations of the future
will be confined to small, isolated protected areas, so that
numbers are too low to conserve original genetic diversity.
Without active and strengthened effort to conserve catson private lands, or to maintain habitat corridors which
restore connectivity to populations isolated by habitatchange, this may indeed come to pass.
Of all the cats, the problem of too-small, isolated populations is most advanced for the tiger, and it is probably
for this species that the resources of the zoo community
will be called upon first. Plans to develop a genome
resource bank for the tiger are timely (Wildt et al. 1993b).
Tigers are well-represented and well-managed in captivity,
and those responsible for their oversight are guided by aninteractive strategic plan which has as its “paramount purpose... contributing to the survival and recovery of all distinct taxa in the wild,” and recognizes that “captive
populations are a support, not a substitute, for wild populations” (Tilson et al. 1992).
However, tigers are seriously threatened by commercial
poaching, and it is of top priority to protect individual populations effectively and actively monitor their status. “Reinvigorating” these small populations can only take placewhen the poaching threat has been brought under control
so that populations are relatively stable and their demography, dynamics, and genetics can be assessed.
Therefore, while it is important to maintain viable captive populations and continue to refine assisted reproduction in captive animals, neither is likely to be applied tomade by the IUCNESC Conservation Breeding Specialist
Group (CBSG) through their Population and Habitat
Viability Analysis (PHVA) workshops-discussedin
Chapter 3-has been particularly significant. The generalpublic, particularly in the developed countries, is becoming more aware of the need to conserve wildlife and, aszoos compete for people’s leisure time with a growing
number of other options, it is likely that zoos will increasingly seek to demonstrate their commitment to conservation. Four areas where zoos have the potential to makesignificant contributions are reviewed.

Maintaining
Populations

Viable Captiveof Rare Species

Captive breeding is a primary area of zoo expertise, andthe maintenance of genetically viable populations of rareand endangered species has been a major emphasis since
the early 1980s (Seal and Foose 1986, Hutchins and Wiese
199 1, IUDZGKBSG
1993). In terms of species conservation, it is important to maintain viable populations ofrare species in captivity for the following two purposes(IUCN 1987):
1. As an insurance population for possible reintroduction
should the species become extinct in the wild or overpart of its range;
2. As a source of new genetic material to infuse diversity
into depleted wild populations.
The status of captive cat populations and their self-sustainability has been reviewed in this chapter. With the
exception of the big cats, captive populations are biased
toward more common species. The small cats are relatively poorly represented, particularly the rare Categoryl-3 species, and most regional populations are not self-sustaining. This situation is unlikely to change until research
into the status and ecology of these small cats, as recommended by this Action Plan, is carried out: it is inadvisable
to remove individuals from wild populations of these
species unless basic studies have been done. There may be
more individuals of these species held in captivity, some inprivate collections, than are known at present (P. Quillen,
pers. comm.).
The North American and AustralasianRegional
Collection Plans recommend that zoos reduce their holdings of the more common and easily reproduced species
to free captive space for those that are less common but,for the near future, acquisition of new founders or unrelated animals depends upon international cooperation with
reputable zoos in the range states. Years of work lie ahead
before all of the rarer species of the family Felidae are
maintained in viable captive populations.

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Research
Zoos are increasingly looking to wild populations to help
them better manage their captive populations, with thecheetah being a good example among the cats. Becauseof problems with poor reproduction and susceptibility todisease in captivity, the North American Cheetah Species
Survival Plan (SSP) is studying wild behaviors to help
design more appropriate captive habitat for cheetahs, andis collecting information on incidence of disease in wild
populations (Caro 1993, Laurenson 1993; L. MarkerKraus, L. Munson, pers. comm.).
Conversely, field biologists should increase cooperation with the zoo community to improve the conservation
management of wild populations. For example, there is a
wealth of knowledge among breeders about biological andreproductive parameters of cat species, which can provide
baseline data to aid understanding of the dynamics of wild
populations.
For most cats, the data in the Species
Accounts under the heading Biology is derived from captive animals, and is just an indication of the volume ofphysiological data which could be collected.

can promote

publicsupportfor wildlife

conservation

Chapter

5. Cats in Captivity

Zoo animals have provided the foundation for thedevelopment of classic descriptive biological sciences,including anatomy, morphology, taxonomy, classification,
study of locomotion, nutrition, and other aspects of behavior. Captive studies, particularly the work of Leyhausen(1979), have greatly advanced understanding of cat behavior. Newer scientific disciplines also benefit from studies
undertaken on captive animals, including small population
biology, genetic and molecular studies, wildlife veterinary medicine, pathology, parasitology and bio-energetits (Hutchins etal. 1991, IUDZGKBSG
1993, S. O’Brien
pers. comm.). Of special importance to field biologists,
methods of individual identification can be developed
from and tested on captive animals (e.g., Miththapala et al.
1989).

wild cat conservation in the near future. In the meantime,zoos can contribute more immediately by increasing theirrole in the areas discussed below, most importantly public
education and direct financial support.

Zoos

Issues.

Public Education
The World Zoo Conservation Strategy (IUDZGKBSG
1993) estimates that the world’s 1,100 organized zoos
annually receive at least 600 million visitors, over 10 percent of the entire world population. Zoo visitors include
people of all ages from a broad spectrum of social, ethnic,
cultural and educational backgrounds, all of whom have
one thing in common: an interest in animals. In addition tocasual visitors, many formal education groups visit zoos,
often consisting of children, but also including university

with displaysof animals.

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wildlife conservation
could become more directly
involved by helping to fund high priority in situ conservation projects. As pointed out above, special exhibits can becreated around these projects which can serve as a draw
to visitors.
Traditionally, zoos have invested comparatively little
in field conservation (Hutchins and Wiese 199 1), althoughsome, such as the Jersey Wildlife Trust, have been outstanding in their efforts. In addition, several large publiczoos support semi-independent zoological societies whichplay a major role in wildlife conservation (e.g. the Frankfurt
and New York Zoological Societies and the ZoologicalSociety of London). Aside from the work of these organizations, we are aware of very few examples of zoo funding for field projects which specifically benefit cats.
The Tiger Global Animal Survival Plan (Tilson et al.
1992) recommends that participating zoos cooperate toprovide $925,000 a year for 10 years to irt situ tiger conservation, especially through “Adopt-A-Park”
programs.
The GASP points out that there are over 325 “hard currency” zoos with tigers, and the level of financial commitment sought represents an average of U.S. $2,770 per
ins titution, or j ust over $1,000 per captive tiger.
It would be possible for individual zoos collecti vely tomake a substantial contribution to wild cat conservation if
a small annual donation were paid into a fund for the
implementation of priority projects along the lines of thoserecommended by this Action Plan. By keeping the per animal donation low, smaller zoos would not be disproportionately burdened and it is likely that zoos would not findthe request unreasonable. In return, zoos could expect toreceive progress reports so that appropriate displays could
be created around their involvement. Rather than have towork out cat conservation projects on an individual, ad hoc
basis, zoos could legitimately publicize direct financial
support for a number of projects involving various species.
Such a level of funding support from the zoos of theworld is entirely appropria te -a “conservation duty”and would not be so onerous as to rule out support forother projects. Until such time that interacti ve management of wild an.d capti ve populations becomes an effective
and widely used tool, zoos could make a substantial,
immediate and urgently needed contribution by increasing their direct support of conservation of wild cat populations (Action Plan Project 14).

and pre-university students.
The live animals exhibited by zoos can give meaning
and immediacy to conservation education far beyond that
achieved by film or print. Many zoos now have attractive
settings for big cats, but much could be done to improve thedisplays of lesser cats to promote wider public interest.
The World Zoo Conservation Strategy emphasizes thatconservation education is a key element in zoo education,
which in its most basic form consists of information about
the threatened status of an exhibited animal and an explanation of the causal factors. Animal exhibits, however, canbe used to explain a variety of biological subjects, ranging from species-specific ecological and behavioral adaptations to broader themes such as ecological balance,
evolutionary radiations and biological diversity. Creative
exhibits and educational material at zoos are an invaluable resource for the cause of wildlife conservation, withenormous and still under-realized potential to stimulate in
the general public an abiding interest in wildlife and concern for its preservation.
Wildlife conservation is of international importance,
but depends on local action. The zoo forum is an appropriate one for educating people about various approaches
to conservation around the world, and for helping toengender in the public a more sophisticated grasp of theissues involved. To this end, there should be greater cooperation between field conservationists and zoo educators.
In view of the public’s fascination with cats, it would be
ideal for zoos to display educational material highlighting
key projects and the work of organizations concerned withthe conservation of wild cats. The Cat Specialist Group is
well-positioned to play an advisory role in setting up suchdisplays and exhibits. In this way, guidance would be provided to those visitors wanting to become more actively
involved. This pertains especially to pre-university students; the importance of recruiting more field conserva tion
specialists has been emphasized in Chapter 3.

Helping
Wildlife

to Pay the Costs ofConservation

Conservation benefi ts arising from zoo research and education of visitors are important, but are largely diffuse andindirect. Zoos seeking to play active roles in the field of262

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Chapter

6. Reintroduction

Part II
Major Issues in Cat Conservation

Chapter 6
Reintroduction
reintroductions have taken place in developed countries, orin those African countries with well-funded wildlife
departments. Few projects, however, have been carefullyplanned and executed, although there are exceptions.
Three examples of well-planned reintroductions are discussed below.
In Germany’s state of Bavaria, 129 wild-caught andcaptive-bred European wildcats (75 males and 54 females)
were released over a period of five years (1984- 1989) atthree sites in state-owned forest. Planning and collectionof the animals to be reintroduced began several years prior
to the releases. After the releases, the animals were monitored by radio-telemetry and by distribution of questionnaires to local residents, workers, and hikers. There washigh road-kill in the first weeks following release, and survival was estimated preliminarily to be quite low, of theorder of 20-30% (Biittner and Wore1 1990). Still, there
was evidence of reproduction at all three release sites.
In the Adirondack mountains of northern New Yorkstate, 83 Canada lynx (48 males and 35 females), wildcaught in the Canadian Yukon, were released over three
winters from 1988- 1990. Twenty-three lynxes had died byearly 1992: 12 were hit by cars, five were accidentally
shot, and six died from miscellaneous causes, including ayoung lynx killed by an adult male. Three lynx raided
livestock pens. Some lynx migrated far from the releasesites, but human-related mortality was low within New
York state, due in part to substantial pre-release publicity
and public education. While there is still no direct evidence of breeding, there have been unverified sightings ofkittens (Brocke and Gustafson 1992).
In Georgia, 32 bobcats (15 males and 17 females),wild-caught on the coastal plain, were reintroduced in
1988- 1989 to Cumberland Island, near the southern
Georgia coast. The island is largely undeveloped, and preydensities were found to be high. All of the reintroduced
bobcats were radio-collared; blood samples were taken
prior to release; and the project received widespread publicity through the media. Several graduate students did
thesis work around different aspects of the reintroduction,

Introduction
The Species Survival Commission of the IUCN includes
the Reintroduction
Specialist Group. The group hasdrafted guidelines for reintroduction projects, which areavailable from the SSC office at IUCN headquarters in
Gland, Switzerland.
The umbrella term “reintroduction”
includes several
different concepts as defined below:
1. Reintroduction: in strict terms, an attempt to establish
a species, subspecies, or race in an area which was once
part of its historical range, but from which it has
become extinct. Re-establishment is a synonym, but
implies that reintroduction has been successful, something that may require considerable time to assess.
??~nsZocation: movement of individuals or populations
from one part of their range to another.
Reinforcement/Supplementation:
addition of individuals to an existing population of the same species, subspecies, or race.
Conservation/Benign introduction: an attempt to establish a species, for the purpose of conservation, outside
its recorded distribution, but within an appropriate %habitat and eco-geographical area.
The following sections review recent felid reintroductionsof these various types. In concluding, the lessons learned
from these projects are assessed, and the role reintroduction has to play in wild cat conservation in the 1990s is
evaluated.

Reintroductions
Several cat species have been reintroduced to parts of theirrange from which they had become extinct. All of these

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Chapter

6. Reintroduction

troduction schemes could not be regarded as a priority
strategy for conservation of the European wildcat.
Attention should instead focus on conservation of existing populations.
An example of the sort of unpredictable demographic,
environmental and genetic changes that can affect small
reintroduced populations is a series of events which have
befallen the lynx population reintroduced in Switzerland
over 20 years ago. Mortality has been so high, primarily
due to illegal killing and road deaths, that the radiationand growth of the population has stopped. In the Jura
Mountains sub-population, all but one resident male havebeen killed. That male has expanded his home range tocover the home ranges of six females, thereby disproportionately increasing his genetic representation in the population. The lack of other competing breeding males isrelated to poor cub survival. Observations and estimates
showed that a maximum of 3-6 of 22 young in the Jura
reached their third year, and all of them were female(Breitenmoser et al. 1994).
Another example is the lion population of the
HluhluweKJmfolozi
reserve complex in South Africa. A
nomadic male took up residence in the park in 1958, andseven years later two adult females and two young cubs
were released. The population today is thus descended
from a very small gene pool: the two females came fromthe same pride, and it is presumed that they were related,while the original cubs were likely to have been killed by
the male (Anderson 198 1, 1992). In 1993, male lions fromthe reserve were found to have very low sperm quality(M. Briggs in litt. 1993).
Successful reintroduction relies upon careful planningand preparation, which requires both time and money. If
this is not to be wasted, it is in the interest of management
authorities to monitor reintroduced populations for at least
several decades following release, and to be prepared totake active measures if necessary to prevent reextinction.
Another problem which will affect reintroductions
where cats have been absent for long periods is the impact
of predators on inexperienced prey populations, both wildand domestic. When released in Switzerland, lynx had
been extinct for at least 100 years, and reintroduced animals killed considerable numbers of roe deer and chamois(Breitenmoser and Haller 1993). Reintroduced lynx also
killed domesticated sheep. From 1984-1988, 135 attacks
were reported in an area of France adjoining the Swiss
Jura, and in 1989,426 attacks were reported within an areaof only 50 km2 (Herrenschmidt and Vandel 1989, Anon.
1990d). Breitenmoser (1983) found that sheep made up
20% of lynx kills. Livestock depredation in France andSwitzerland arouses considerable controversy and publicantagonism despite compensation schemes. Similarly,
cheetahs released in several southern African reserves had
to be removed because of their high levels of predation on

including feeding ecology and effects of reintroduced catson prey populations (Baker 1991), population dynamics
and evaluation of census methods (Diefenbach 1992), andsocial organization and habitat use (W. James in prep.).
Most animals were recaptured for examination after the
first year following release, and they were found to haveincreased their weight by an average of 12.3% (Diefenbath 1992). There was also evidence of breeding in thefirst year, with four dens and 10 kittens found (Baker
199 1). Diefenbach (1992) estimated adult survival rate at
93% and juvenile (~1 year) survival at 35%. The population more than doubled following the first year of release,and reproduction apparently declined after three years as
density increased.
Other reintroductions have been less carefully planned.
These include servals, cheetahs, and lions in various South
African reserves (van Aarde and Skinner 1986, Anderson
1992); wildcats and Eurasian lynx in several European
countries (Breitenmoser and Breitenmoser-Wursten
1990,
Stahl and Artois 1991); and pumas in several southern andwestern American states (Jordan 199 1, Homocker 1992).
Most of these efforts involved the release of fewer than
20 animals, and in some cases fewer than five. Only therelease of lynx in Switzerland has involved rigorous follow-up monitoring. Although monitoring only began 10
years after the releases, it has continued for 10 years andis now the longest running follow-up radiotelemetry studyof a reintroduced cat population.
How successful were these reintroductions‘? It depends
upon the time frame used to measure success. Many reintroduced cat populations are surviving today, and some,like the bobcats of Cumberland Island or Eurasian lynx
reintroduced to Slovenia, have rapidly increased in number
and-inthe case of Slovenia-haveconsiderably
expanded their range (Cop 1992). However, long-term
persistence of any of the populations cannot be assured.
Small populations are vulnerable to severe fluctuations
caused by environmental
or demographic
factors.
Diefenbach ( 1992) developed population viability models
for the Cumberland Island bobcats and, based solely on
demographic factors, estimated the median persistence
time of the population at only 65 years. The island (only
80 km2 in size) may be too small to support a viable bobcat population, even with supplemental releases to minimize inbreeding.
In a review of the wildcat reintroduction in Bavaria,
Stahl and Artois (199 1) commented that even with a project of that scope, which had existed for 10 years, there wasno guarantee that viable wildcat populations had become
established in any of the release sites. They added that
greater care was needed before launching major reintroduction programs.
They suggested that the risks ofhybridization with domestic cats and the high degree of
threat resulting from habitat modification meant that rein-

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small populations of antelopes in fenced reserves whichwere not previously exposed to to large predators (Pettifer
1981, van Dyk 1991, Anderson 1992).
Cat reintroduction projects can expect predators to havea strong initial impact on prey populations. Although preyanimals can regain their escape instincts fairly rapidly(Breitenmoser and Haller 1993), the consequences could
be more serious inside small fenced areas, such as wildlife
reserves.
Reintroduced wild-caught cats generally survive better
than captive-bred animals. This is because animals rearedin captivity have not developed foraging skills, and have little fear of humans. Captive-raised cheetahs in South
Africa, for example, repeatedly raided the chicken housesof people living on the game farm where they werereleased, even in daylight with people sitting about (andthrowing stones) (van Dyk 1991). However, it is possible
to raise captive-bred animals so that prey-capture skills arelearned. The Bavarian wildcat reintroduction project beganwith four wild-caught pairs. When they had bred, themothers and young were kept in enclosures where live prey
was released. After six months of age, the young cats weremoved to acclimatization enclosures near the release site
and fed. They were released shortly thereafter, with supplemental food supplies (Buttner and Wore1 1990).

Translocatedtiger being released
stock at the edge of the park.

in Khao Yai National

Issues.

Chapter

6. Reintroduction

With proper care and precautions, captive animals canfare as well, if not better, than wild-caught cats. Captivebred pumas released in Florida have not only survived andreproduced (Jordan 199 1, 1994), but also appear to be settling down in the release area more readily than translocated wild-caught animals (Belden and McCown 1993).
Even with wild-caught animals, it is recommended thatthey be kept in large enclosures near the release site for ashort period, to allow them to recover from capture andtransport stress, eat, and become acclimatized, rather thanreleasing them directly into the wild (Diefenbach 1992).

Translocations andPopulation Supplementation
Although translocation and population supplementation
are often described as conservation tools of the future,
which will permit intensive genetic management of small
isolated cat populations, experimentation has actually been
underway for decades. These efforts have been local andinformal, and thus documentation is lacking. Documentation is most readily accessible for translocations of big
cats in eastern and southern Africa but, as with most rein-

Park, Thailand,265

in 1988.

It had been taking

live-Part II: Major

Issues.

Chapter

6. Reintroduction

Tranquilized
lions being transported
National Park, Bophuthatswana.

by air from Etosha

in Namibia

for reintroductionin the Pilanesberg

Translocation has often failed, and there are no guidelines for ensuring the success of a translocation attempt.
Some animals establish residence at the release site, andothers migrate long distances. The behavior does notappear to be species-specific-it
may be highly individualized, or depend upon conditions at the release site. As an
example of a successful translocation/population supplementation, the two lionesses introduced into Hluhluwe/
Umfolozi in South Africa (where there was only one resident male at the time) were translocated from Kruger
National Park, more than 100 km away. They were thefounders of a population now estimated at 100 animals(Anderson 198 1, 1992). However, in some cases cats have
moved far away from their release sites (Hamilton 1981),

troductions, few of these attempts have been very systematic, and none have involved long-term monitoring.
Lions, leopards, and cheetahs, typically problem animals, have been translocated and introduced into reserves
where populations were considered to be low. As an
example of more deliberate attempts to increase population size, cheetahs were released into South Africa’s
Kruger and Kalahari Gemsbok National Parks, and into
Namibia’s Etosha National Park, in the 1970s (du Preez
1970, Anderson 1992). Although some of the animals
were marked with ear tags, few were subsequently
resighted, and resighting effort was not systematic, so thatthe effects, both immediate and long-term, of population
supplementation are unknown.

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even if there were no resident animals present to drive
them off (Gossow and Hdnsig-Erlenburg
1986, Jordan
199 1). One translocated leopard in South Africa travelled
more than 540 km back to his original home range area
(Jewel1 1982).
The importance of better understanding of the complexities of injecting an “outsider” into an existing felidsociety is illustrated by an attempt to translocate a tiger
which killed a woman at the edge of the Indian Sundarbans
mangrove delta. It was released in the interior of the
Sundarbans Tiger Reserve, but was killed by another,larger tiger only 20 meters from the transfer cage, to which
it had been seen to return a short time before (Seidensticker
et al. 1976). In a second translocation in the Sundarbans, atiger which had taken livestock was released elsewhere,
swam a creek, and disappeared immediately amongst the
mangroves (Ghosh 1988). Its ultimate fate is unknown.
While knowledge of the results of translocations andsupplementations remains generally obscure and localized,
there are a few well-publicized, even notorious, examples.
These translocations were carried out for the welfare ofindividual animals, rather than strictly in the interest of
population conservation.
Captive-raised animals were“returned to freedom in the wild” by release in parks holding resident populations. Examples include the release oflions, leopards, and a cheetah by Joy and George Adamson(Adamson 1960,1969,1986), and the release of a tigress
and two leopards in India’s Dudhwa National Park by
Arjan Singh (1981,1984). These projects were marred byinjuries and deaths to humans, probably due to the cats’
familiarity with people and lack of natural caution.
In the Indian case, an outbreak of tiger attacks on people
soon after Singh’s tigress, Tara, went wild in 1978 led to
public accusations that she was responsible, a cry taken up
by politicians and used to attack the cause of wildlife conservation.
Leading Indian tiger specialists, however,believe that the attacks on people, which continued into
1993, were an unfortunate coincidence, and that Tara hadprobably been killed by a resident tiger. Singh ( 198 1), on
the other hand, is convinced that Tara was integrated into
the local tiger population and subsequently bred. The affair
is an illustration of the political perils of reintroductions.
One of the main scientific concerns involved in translocation and population supplementation is that introduced
animals should not introduce “foreign” genes into the resident population. A feature of the Adamson and Singhreleases, and of a later release of lions from Kenya into
Botswana (Anon. 1991a, b, 1993e) was that the animals
were not of the local race and, moreover, that some were
genetic “cocktails” due to cross-breeding with other subspecies or generics in captivity. Wirth (1990) quotes thelate Sir Peter Scott (then chairman of the IUCN Species
Survival Commission) expressing concern that the tigress
released by Singh was a hybrid of two races, neither ofIssues.

Chapter

6. Reintroduction

them the same as the Dudhwa population. Scott added that
if she bred it would “make a further genetic cocktail, andfrom a scientific and conservationist point of view thiswould be deplorable.”
However, while the issue is obviously important, clearcut guidelines will be slow to emerge. How significant
were genetic or adaptive differences between, for example,
the Kenya lions and the resident Botswana lions? There
was no attempt to evaluate this before release. However,preliminary genetic and morphological analysis of African
leopards indicates that all these leopards, despite the very
different habitats (sand desert, tropical rain forest) inwhich they are found, could be considered to belong to asingle subspecies (Miththapala 1992). Can the same besaid for lions? Analysis of the mitochondrial DNA of several populations of east African antelope species revealed
great interspecific variation in the amount of genetic differentiation between populations (P. Kat in Zitt. 1993).
Furthermore, in some cases the introduction of new
genetic material may be desirable. O’Brien et al. (1990)
compared the genetics of two isolated populations of
Florida panther. One population had been supplemented(not officially) by the release of hybrid North and SouthAmerican pumas, some of which had bred so that theirgenes entered the population’s gene pool. The genetic
diversity of this population was substantially greater thanthat of the isolated pure population, leading to the conclusion that hybridization can be beneficial for small inbred
populations (CBSG 1991).
Another aspect of population supplementation whichneeds further examination before it can be recommended
as a conservation tool is how introduced animals are
accepted by the resident population. Under what circumstances can introduced cats be expected to establish a territory and be incorporated into the breeding population?
Under what circumstances are immigrants likely to be
killed? Are the chances of success higher with females
than with males? These questions should all be investigated before population supplementation is seriously considered as a high-tech solution to the genetic problems
faced by small wild populations.
In the planning of any felid translocations or reintroductions, the risk of accidental transmission of disease into
unaffected populations via released animals must be carefully assessed. Woodford and Rossiter (1993) recommendthat veterinary involvement in reintroduction projects
should begin as early as possible. They point out instancesof inadequate disease risk assessment resulting in expensive failures, and the introduction of destructive pathogens
into resident wildlife populations by captive-bred andwild-caught animals. Infectious diseases may have shortor long-term effects on population size and viability byaffecting rates and patterns of mortality or reproduction.
The assessment procedure should address all the infec-

267

Part II: Major

Issues.

Chapter

6. Reintroduction

population density are probably ideal, but such areas are
now few and far between (see Chapter 1). Human persecution is a major cause of cat extirpation, and if cats are to
be reintroduced to settled areas, substantial pre-releasepublicity and communication should be undertaken. Reintroduced cats may roam a great distance from the releasearea, enter settled areas, and kill livestock or people. As
pointed out by Moore (1992), because carnivores arewidely viewed as “villains” threatening livestock and people, the most important factors in the survival of reintroduced carnivores are often socio-political.

tious agents to which the animals to be moved might havebeen exposed, and could carry, and which might affectconspecifics or other species at the destination. In this context the list of diseases to be considered may be longer thanthat in Table 5 in the chapter on Research.
For some infections, tests are available, which, if negative, provide confidence that the animal is not a carrier.
For others (e.g., tuberculosis), testing may not be straightforward or reliable, or there may be no test available (e.g.,spongiform encephalopathy). It should be borne in mind
that several diseases of felines (e.g., feline immunodeficiency virus disease and feline spongiform encephalopathy) have been recognized only very recentlv and it is notunlikely that other, as yet unrecognized, infections may
occur in cats.
The details of the appropriate protocols for quarantine
and testing procedures to be undertaken prior to shi pment
or release will depend upon the species involved andknowledge of the infectious disease status of the donor andrecipient populations, and of other species with which they
are in contact via a food chain or other routes.
Predatory training (of captive animals), genetic selection, disease screening and quarantine, immobilization,transportation, and release add up to considerable expense,
which will be wasted if introduced animals fail to survive
and breed. It is unfortunate that the results of introducing
new animals were not more closely monitored in previous
attempts, but there is likely to be opportunity in the future,
as translocation remains an important management tool
on a local scale. For instance, when faced with limited
options regarding how to handle a rare big cat which is
causing problems with people and livestock, translocation
is often considered an attractive alternative to elimination(see Chapter 2).
Because these issues are likely to take on greater importance to cat conservation in the next century, it is notargued here that experimentation with translocation andpopulation supplementation cease, only that they be more
carefully undertaken and monitored.

Summary of Problems
Reintroducing
Cats

Release Areas with Adequate
Suitable Food Supply

Prior to reintroduction, prey species in the release area
should be censused, and their numbers evaluated to see
roughly how many cats they are capable of sustaining.
Predation rates can be calculated from studies of kill rate in
the wild, or from maintenance energy rates in captivity,
coupled with what is known of the species ecology-e.g.,
females with young have higher energy requirements, andwhile most cats will utilize nearly all of a large kill,remaining near the carcass until it is substantially consumed, cheetahs require fresh meat and may kill large animals

Prey
been
such
than
dyna
declines due to prey inexperience with predators.
For supplementation of depleted cat populations, managers should ensure that prey decline was not among thefactors leading to the decline of the cats.

Source of Animals

For reintroduction,“wilderness”

to be Reintroduced

As Yalden (1993) pointed out, there is general agreementthat reintroduced animals should be taxonomically andgenetically as close as possible to the former population,
yet an objective definition of “suitability” has not yet beendeveloped, and cannot realistically be expected to emerge
for some time. There needs to be substantially more
research on intraspecific diversity before good general
guidelines can be formulated. For the present, according
to local priorities and sensibilities, it is up to local managers to weigh the ecological significance of an ecosystem having (a) no cats where they formerly existed or (b)cats of potentially different genetic makeup than those thatformerlv existed.
The same consideration applies to introducing animals

withThis review, in addition to those of others (Wemmer andSunquist 1988, Diefenbach 1992, Yalden 1993), points toa set of key considerations which should be undertaken
before reintroduction of cats, especially the big cats, iscontemplated seriously.

Release Areas Acceptableandto People
areas with low human268Part II: Major

of potentially different genetic makeup to resident, albeit
severely reduced, populations. The draft guidelines of the
IUCN Reintroduction Specialist Group advise against it.
However, managers may some day face a real dilemma:
is it better to let a population go extinct, or become highly
in bred, rather than “contaminate” it?
These questions highlight the importance of linking
field studies and genetic/morphological analysis to shed
more light on the significance of ecological adaptation andgenetic diversity within cat species. Answers to thesequestions are likely to be species-specific, and will require
substantial research and field collection effort to obtain.

DiseaseRisks

Precautions are essential against introduction of disease
to wild populations by reintroduced animals. Assessmentof all the infectious agents to whi ch animals to be reintroduced have been exposed should be carried out and quarantine requirements respected. The advice of the IUCN/SSC Veterinary Specialist Group should be sought.

Developmentof a reintroduced

lynx population

is being studied

Issues.

Chapter

6. Reintroduction

Costs of Reintroduction
Reintroduction can be very expensive, especially if captive
animals are involved which have to receive acclimatization and predatory training. For example, an ambitious
plan has been developed to breed wild-caught Florida panthers in captivity for eventual population supplementation.
The plan estimates costs of building a captive facility,
developing reproductive technology, and genetic analysis
at over U.S. $500,000 (Seal et al. 1989). Wemmer andSunquist (1988) estimate the capital cost of constructing asnow leopard propagation and holding facility in a snow
leopard range state at over $200,000, with annual maintenance costs of nearly $20,000.
Even if wild-caught animals are used, costs of publicity and communication, capture, interim feeding and maintenance, animal evaluation (clinical examination, blood
samples), equipment (telemetry, vehicles) and follow-up
monitoring (personnel support) can be cumulatively high.
It is also very important that project budgets take the costsof long-term future monitoring into account, particularly if
initial project expenses are high.

in the Swiss

Jura Mountains.

Part II: Major

Issues.

Chapter

6. Reintroduction

to the proximity of a large, potentially dangerous predator, will be living around, if not actually in, any reintroduction site in India.
However, on a regional or local scale, reintroduction,translocation and population supplementation can be very
important tools. For example, although the viability ofsome of the populations is questionable, much of the westem range of the European wildcat and Eurasian lynx wascolonized in the last few decades by reintroduced animals(Breitenmoser and Breitenmoser-Wiirsten
1990, Stahl andArtois 199 1). While these species are not globally endangered, their re-appearance in western Europe is of great
regional significance. Similarly, while the reintroductionof Canada lynx to northeastern New York is not of great
advantage to the survival of the species, it is important for
the “Lower 48” United States, where lynx populations are
small, threatened, and widely separated.
All parties interested in reintroducing cats are urged tocontact both the Cat and the Reintroduction Specialist
Groups. The considerations enumerated above should betaken into account before any animals are actually
released. Monitoring of recently reintroduced populations
should continue, and future research should evaluate theaspects of predator reintroduction which remain in question, including:Conclusions: What is theRole of Reintroduction in
Cat Conservation?
If highest priority for reintroduction were to go to thosespecies which have disappeared from substantial portionsof their former range, the list for cats would include the following: lion, tiger, leopard, and cheetah (north Africa andparts of Asia); jaguar (southern U.S. and north-central
Argentina); and puma (eastern North America). But direct
human persecution and depletion of the prey base were themajor causes of decline. In the case of the north Africanand southwest Asian cats, antelope populations, which
would provide prey, are still in a seriously depleted condition (East 1992a, b), and it is unlikely that local people
are ready to support reintroduction.
It is also questionable whether the current stock of captive animals is genetically representative of extinct populations or, if they are
not, whether they are suitable substitutes. ln the case of thejaguar, the northern and southern parts of its former range
are now heavily settled, and hardly present ideal conditions
for reintroduction. Only in the case of the eastern cougar
are conditions suitable for a widespread reintroduction
effort (particularly in northeastern North America), andthen the sporadic sightings of wild animals (indicating that
there are perhaps survivors) and the lack of authentic eastem cougars in captivity are stumbling blocks.
In short, for the 1990s at least, it is unlikely that reintroduction will play a role which is significant on a global
scale, even for those cats which need it most. However,work which pertains to the reintroduction of these catsshould continue. For example, the puma reintroduction
in northern Florida and translocations elsewhere in the
United States (see Species Account) should continue tobe monitored and lessons drawn. The ecology of jaguars
living near settled areas, and management solutions tominimize livestock depredation, should continue to bestudied. For the north African and southwest Asian cats,
the Iranian, Egyptian, and Algerian cheetahs (the beststock for future reintroductions elsewhere in the region)
should be protected and their ecology studied.
Reintroduction of Asiatic lions to a site in the former
range continues to be under active consideration becauseof their current existence as a single, isolated, high-density and low genetic diversity population in an Indian
reserve. Johnsingh et ul. (199 l), Ravi Chellam andJohnsingh ( 1993b), and Walker (1994) favor the establishment of a second population (see Species Account).
However, the socio-political aspects need careful consideration, since people, including livestock owners, unused

1. The conditions under which translocation of individuals
to resident populations is successful;
2. How predatory training of captive-raised
most efficiently be accomplished;

animals can

3. How animals differ genetically and morphologicallydifferent parts of their ranges, and;in

4. How people can be encouraged to tolerate or even facilitate the return of a potentially dangerous or destructive felid predator.
The final point needs to be stressed because of the impact
carnivores can have on human communities, which arealmost always involved. The comments of Jo Gipps of
the Zoological Society of London in a preface to a symposium on reintroductions (Gipps 1991) are especiallyrelevant as far as big cats are concerned: “However good
our understanding of the biology of a species and its habitat, conservation projects are almost inevitably doomed to
failure unless the dynamics of the human relationships arewell understood by everyone involved, and unless thoserelationships are based on mutual respect and understanding of each other’s problems.”

270Part III: Cat Action

PlanPart IllAn Action Plan for Cat
Conservation in the 1990s

I
II
III
IV
V

Introduction
The following projects are priorities for wild cat conservation over the coming decade. They are presented in a
simple summary form. The projects focus on the most vulnerable species, and are organized into two sections.
Section 1 consists of general projects which address major
issues in cat conservation, as defined in Part II of this document. The second, larger group of projects is laid out inthe same order as the Species Accounts. To locate projects
for certain species or regions, see the list of projects which
follows. Although these projects are numerous, they do
not address all species, nor all the general recommendations made for future cat work in Part II.
There are two types of projects included in the Action
Plan. The first type consists of existing projects whichhave received varying amounts of their budgeted funding.
For these projects, a contact address is provided for donors
and other interested parties. They are distinguished from thesecond group by the placement of an asterisk * after the title.
The second type of project consists of those proposedby Cat Specialist Group members. These projects needfunding and, in many cases, workers.
Donors and otherinterested parties should contact the Cat Specialist Group
directly for details. The Chairman maintains a databaseof telephone, fax, and e-mail contacts for the executors ofexisting projects, and asks that the Group be informed ofcontacts related to these projects.
The Cat Specialist Group’s contact addresses are: Peter
Jackson, Chairman, IUCN/SSC Cat Specialist Group,1172 Bougy, Switzerland, Tel/Fax +4 1 2 1 808 60 12, email peterjackson@gn.apc.org
or c/o Species Survival
Commission, IUCN-The
World Conservation Union,
1196 Gland, Switzerland, Tel: +41 (22) 999 0001, e-mail:mgd@hq.iucn.ch; Kristin Nowell, 2520-4. 41st St. NW.
Washington DC 20007, U.S.A.
Projects are categorized as follows in terms of approximate annual budget (all figures in 1994 U.S. $ ):$10,000 or less
$1 o,ooo-$20,000$20,000-$50,000$50,000-$100,000
over $100,000* = Ongoing Project

List of Priority
I. General

Projects

Topics

Implementation of the Cat Action Plan
1. Establishment of the Cat Conservation Data Center
Habitat loss and fragmentation
2. Response of a felid community to logging of tropical Asian rain forest
3.

Acquisition of map databases for overlay of cat distribution survey data

4.

Identification of potential protected areas for conservation of biodiversity in the Indian Himalayas*

Management of big cats near people
L5 .
Global survey of methods and techniques to minimize the impact of livestock losses to cats
6.

Support for the National Center for Research, Management, and Conservation of Predators in Brazil*

Research
7. A guide to census procedures for cat populations

271

8.

A workshop to define minimum viable population
sizes for cat species

9.

A fund for field collection and processing of biological samples for genetic, morphological, and clinicalanalysis

Part III: Cat ActionPlan

10. Systematics
problems*of the genus
FeZis and hybridization.

30. Support for the Serengeti Lion Project*
31. Support for the N!aiuh Project in the Namibian
Kalahari*

11. A survey of disease in wild cat populations*

32. Resolving lion and livestock conflicts in west Africa

Trade
12. Understanding the market for tiger bone medicines*

Serval (Category 3)
No specific projects suggested, but see related projects.

13. The leopard cat in China: ecology and management
for sustainable utilization*

Leopard (Category 4A)
33. Evaluation of the effects of sport hunting on leopard
population dynamics

Cats in Captivity
14. Establish a zoo-based fund for field conservation ofwild cats

34. Leopard ecologv and density in tropical African rain
forest*“d

15. Evaluation of subspeciation and establishment ofregional captive breeding programs for the wildcat*

Caracal (Category 4)
No specific projects suggested but see related projects.

Reintroduction
16. Long-term monitoring of the reintroduced Eurasianlynx population in Switzerland*

African wildcat (Category 5)
No specific projects suggested but see related projects.

17. Population dynamics of a reintroduced bobcat population in a small isolated habitat block*North Africa and Southwest

Il. Species

Projects

Sub-SaharanAfricaAsia

Asiatic lion (Category 1)
35. Establishment of a second population of Asiatic lions
in India*Black-footed cat (Category 1)
18. Natural history of the black-footed cat*

Cheetah (Category 1A)
36. Conservation of the Asian cheetah in Iran

19. Distribution

37. Natural history, genetics, and conservation status of
the cheetah in the Saharan regionof the black-footed cat

African golden cat (Category 1)
20. Natural history of the African golden cat*

Serval (Category 2A)
38. Survey for the leopard and the serval in the Atlas
Mountains of Morocco*

2 1. Distribution of the African golden cat
Cheetah (Category 2A)
22. Support for the Cheetah Conservation Fund*
23. Factors limiting cheetah populations
tected areas*

Leopard (Category 3A)
39. Distribution and conservation status of the leopard
in north Africa and southwest Asia*

outside pro-

40. Support for the Arabian Leopard Trust*

24. Support for the Serengeti Cheetah Project*

Sand cat (Category 4)
41. Distribution of the sand cat

25. African cheetah status survey*
26. Distribution,
status, and captive breeding of thecheetah in Zimbabwe*

See also related project under manul.

27. Cheetah census and disease surveillance in Kenya*

Caracal (Category 4A)
42. Natural history, distribution,
cal in India*

Lion (Category 2A)
28. Pan-African lion survey*

and status of the cara-

Jungle cat (Category 5)
No specific projects suggested, but see related projects.

29. Predator-prey relationship between lions and large
ungulates in Kruger National Park*

272

Part III: Cat Action

Plan

Wildcat (Category 5)
43. Natural history, status, and captive breeding of thewildcat in Israel*

Rusty-spotted cat (Category 2)
63. Natural history and distribution of the rusty-spottedcat

TropicalFishing cat (Category 2)
64. Natural history of the fishing cat*Asia

65. Distribution of the fishing cat

Tiger (Category 1A)
44. Protection of the Amur tiger and its habitat in the
Russian Far East*

Marbled cat (Category 2)
66. Natural history of the marbled cat

45. Behavior and ecology of the Amur tiger*

See also related project under clouded leopard.

46. Conservation of the South China tiger*
47.

Leopard (Category 3A)
67. Support for the Far Eastern Leopard Fund*

Support for Global Tiger Patrol*

48. Improving the reliability of tiger censuses in India*

68. Leopard ecology

49. Distribution, abundance, and ecological and conservation status of the tiger in India*
50.

52.

Jungle cat (Category 4)
No specific projects suggested, but see related projects.

Conservation of viable tiger populations in India: theeffects of population isolation and movement corridors

51. Support for the Ranthambhore
development Project*Foundation’sin India*

Leopard cat (Category 5)
See project under Trade.

ecoIriomote cat (no ranking)
69. Evaluat .ion of the taxon .omic status of the Iriomotecat*Support for the Tigerhaven Wildlife Trust*

53. Current distribution of tiger populations in east andsoutheast Asia*

70. Conservation of the Iriomote cat*

54. Support for the Chitwan Tiger Project*
55. Development
model*

of a tiger population

Eurasia
Asia sub-region

simulation

56. Genetic evaluation of subspeciation in the tiger*
Bornean bay cat (Category 1)
57. Natural history, distribution,
Bomean bay cat*Snow leopard (Category 1A)
71. Support for the International Snow Leopard Trust*
72. Status and management of the snow leopard in
Tibet*and status of the73. Evaluation of the status of the snow leopard in Russia
and the central Asian republics*

Clouded leopard (Category 2A)
58. Natural history of the clouded leopard*

74. Reducing livestock depredation in the Annapurna
Conservation Area of Nepal: implementation of asnow leopard management plan*

59. Distribution of the clouded leopard and other sympatric small cats*

75. Conservation of snow leopards and their habitat innorthern Pakistan*

Asiatic golden cat (Category 2)
60. Natural history of the Asiatic golden cat

Chinese mountain cat (Category 1)
76. Natural history, distribution,and status of theChinese mountain cat

See also clouded leopard project.
Flat-headed cat (Category 2)
6 1. Natural history of the flat-headed cat
62. Distribution

Manul (Category 2)
77. Natural history of the manulof the flat-headed cat

273

Pat? Ill: Cat Action

Plan

78.

Status of the cheetah, sand cat, and manul inBaluchistan

79.

Status of the manul in the area of the Caspian Sea

Andean mountain cat (Category 1)
92. Natural history, distribution, and status of the Andean
mountain cat
Jaguar (Category 2A)
93. Jaguar and puma depredation of livestock
Pantanal of Mato Grosso, Brazil*

Asiatic wildcat (Category 2)
No specific projects suggested, but see related projects.

94. Ecology of jaguars
Brazilian cerrado*

Eurasian lynx (Category 3)
80. Status survey for the lynx and other small cats in
Xinjiang province, China*

See also related project under Management of Big Cats
Near People.

Iberian lynx (Category 1)
8 1. Implementation of conservation management measures for the Iberian lynx*
82. Applied studies on the conservation
lvnx*

Oncilla (Category 3)
96. Natural history of the oncilla, margay, and ocelotof the Iberian

97. Distribution of the oncillaMargay (Category 3)
See related projects above.

83. Increasing public aware ness of the endangered status of the I berian lynx in Spain*
Eurasian lynx (Category 2)
84. Support for the Status and Conservation
Alpine Lynx Population Project*

Canada lynx (Category 3)
98. Long-term studies of the effects of harvest on Canada
lynx populations*of theGeoffroy’s cat (Category 3)
99. Natural history and distribution of the Geoffroy’s cat
and pampas cat in Argentinian pampas grasslands

85. Spatial organization, population dynamics, and feeding ecology of the reintroduced lynx population in
Slovenia*
86. Preliminary
assessment
Caucasus region*of lynx

statusin thePuma (Category 4A)
100. Support for investigations into the presence of pumas
in eastern North America*

87. Biology and ecology of the lynx in the lowland
primeval forest of the Bialowieza Nature Reserve,
Polandoflynx

populations

10 1. Support for Proyecto Puma in Chilean Patagonia*
102. Projeto Puma: conservation of the puma in southern
Brazil*in

103. Long-term study of puma ecology in southwestern
Alberta, Canada*

European wildcat (Category 3)
89. Status survey and taxonomic evaluation of the wildcats of the Mediterranean island?
90.

in the95. Conservati on of the j aguar in Mexican tropical dry
forest*Europe sub-region

88. Ecology
and status
Scandinavia*

and other carnivores

in theSee also related project under Management of Big Cats
Near People.

Status and distribution of the wildcat in France*
Ocelot (Category 4)
See projects under oncilla.

See also related projects.

Bobcat (Category 4)
104. Distribution and status of the bobcat and puma in the
Mexican Sierras

The Americas
Kodkod (Category 1)
91. Natural history, distribution,kod*

and status of the kod-

Pampas cat (Category 4)105. Evaluation of the taxonomic status of the pampas
cat*

274

Part III: Cat Action

See also related project under Geoffroy’s

Plan

Habitat Loss and Fragmentation

cat.

Jaguarundi (Category 5)
No specific projects suggested, but see related projects.

2.

Priority Projects for Cat
Conservationin the 1990s

Objectives: To evaluate the responses of a felid guild in
previously undisturbed habitat to selective logging; and todevelop recommendations to minimize negative impactsof logging on these species.
Description: This is a pilot project related to conservationof all rain-forest cats. The timber industry is a major partof the economies of most countries with extensive tropical rain forest, and its effects on cat species have not beensystematically investigated. Tropical Asian rain forest is
the most species-rich in terms of cats, and home to several which rank high in vulnerability to extinction. This
study will also lead to increased understanding of how rain
forest cats coexist.
A team of investigators will study the ecology of agroup of sympatric rain forest cat species for l-2 years in
an area scheduled to be logged prior to commencement oflogging activity. Study animals will continue to be monitored throughout the logging process, and for several years
following its completion.
There are three candidate study regions: Sumatra (tiger,
clouded leopard, marbled cat, flat-headed cat, Asiatic
golden cat, fishing cat, leopard cat); Borneo (clouded leopard, Bornean bay cat, flat-headed cat, marbled cat, leopard cat); continental southeast Asia (tiger, leopard, clouded
leopard, Asiatic golden cat, marbled cat, flat-headed cat,fishing cat, leopard cat). It would be useful to conduct atleast two separate studies for comparative purposes.

I. General Topics
Implementation
1.

of the Cat Action

Response of a felid community tologging of tropical Asian rain forest

Plan

Establishment of the Cat
Conservation Data Center

Objective: To assist implementation of the Cat Action Planby establishing a base from which the Cat Specialist Groupcan collect and distribute current conservation-related dataon wild cats. This will facilitate communication between
potential donors of funds and project executants; communication of project results to relevant parties elsewhere;
and the dissemination of information on cat conservation
for the purposes of public education and awareness.
Description: The Cat Specialist Group will establish a permanent center to serve as a coordinating office and clearing-house for data relevant to cats and their conservation.
The center will be central to ensuring that the successes
achieved by priority projects can be duplicated elsewhere,
and mistakes avoided. Through cooperation with theGroup’s research librarian, Gail Foreman, the center will
respond to all requests for papers, contacts, information, orother assistance, with priority going to activities directly
related to Action Plan Projects. Another priority for thecenter will be to coordinate with regional officers the conduct of surveys to improve mapping of cat populations anddatabases on species occurrence in protected areas.
Finally, the center will greatly increase communicationand information exchange between people involved in cat
conservation by continuing to produce the biannual
Specialist Group newsletter, Cat News, and by circulating
copies of papers and reports.

Annual budget: IV-V
Time frame: 5-7 years

3.

Acquisition of map databases foroverlay of cat distribution survey data

Objectives: To improve map-based cat conservation action
planning by: ( 1) evaluation of population fragmentation;(2) identification of important habitat corridors; (3) evaluation of the geographic distribution of inter- and intraspecific diversity; and (4) prioritization of populations forconservation action.

Annual budget: 111
Time frame: Ongoing

Description:
The Cat Conservation Data Center will
acquire Geographic Information System (GIS) map databases, which will be overlaid with field survey data on cat
species presence/absence and population estimates. Map

275

Part III: Cat ActionPlanoverlays include habitat, wilderness, population, settlement and development, protected areas, altitude, precipitation, etc. The map systems will be a great advantage forstrategic planning of surveys, field studies, and conservation action. This project coincides with the plans of the
IUCN Species Survival Commission and the World
Conservation Monitoring Centre to build up a global
species database, including maps, based on data provided
by SSC Specialist Groups.

Time frame: 6 years
Contact: Hemendra Panwar, Director, Wildlife Institute ofIndia, P.O. Box 18, Chandrabani, Dehra Dun 249 001,
Uttar Pradesh, India. Don Hunter, National Biological
Survey, Midcontinent Ecological Science Center, 45 12
McMurry Ave., Fort Collins, CO 805253400, U.S.A.

Managementof Big Cats Near People

Annual budget: I-II
5.

Time frame: Ongoing

Global su rvey of methods and techniques
minimize the impact of livestock losses to

Identification of potential protected areas
for conservation of biodiversity in the Indian
Himalayas*

Objectives: To document a global sample of methods used
to minimize the impact of livestock losses to cats; and todisseminate this information as widely as possible.

Objectives: (1) Using targeted field surveys, Geographic
Information System (GIS) and satellite imagery, to builda model that assesses the general features of high altitude
biodiversity, including the habitat requirements of key
species such as the snow leopard and its prey and; (2) toapply the model to potential reserve sites in the two major
biogeographic regions of the Indian Himalayas.

Description: Although persecution of big cats because of
predation on livestock is a major cause of their extirpation
outside protected areas, there has been no comprehensive
survey of management methods used to control the problem. A global survey of methods to minimize depredation,
sampling a variety of countries, habitats, and species,
would be a useful information source to management
authorities and land owners trying to develop appropriate
solutions to their own depredation problems.
A specialist, working closely with the Cat Specialist
Group, will undertake a global literature review and telephone/mail survey of methods used to minimize livestock
loss to big cats. The survey will be organized according
to representative regional forms of livestock husbandry. A
report will be published in three languages (English,
French, and Spanish), and distributed through the Cat
Specialist Group and other appropriate channels.

4.

Description: Among the more pressing ecological problems facing India’s natural resources is the establishmentof new parks and reserves in the Himalayas-theleastprotected biogeographic region in India, and its richest
region in terms of cat species. Over the next 10 years, thegovernment of India wishes to create a protected area network across the Indian Himalayas, doubling the area coverage. Unfortunately,
this region is among the mostremote and difficult in the world to work in. New methods
are needed that capitalize on limited time in the field forsurveys. This study will develop and test new methods
built around targeted field surveys and augmented bysatellite remote sensing and GIS. Species’ habitat requirements and field survey data will drive spatial models on
biodiversity. The project will attempt to locate key habitat features from satellite imagery and satellite-derived
elevation models. The model will be developed with survey data from existing reserves, and then tested in potentially suitable areas. If the model performs well, themethodology should be applicable throughout the mountains of central Asia. This approach will illuminate thepotential of using models and remote sensing to ( 1) evaluate cat species distribution and status and (2) improvebiodiversity conservation through more strategic planning of protected area network coverage.

Annual budget: III
Time frame: 1-2 years

6.

Support for the National Center for
Research, Management, and Conservationof Predators in Brazil*

Objective: To support a national center of expertise on
predators in Brazil, particularly the big cats, with priority
focus on resolving human/carnivore conflicts.
Description: This is a pilot project relevant to resolving
human/carnivore conflicts in Latin America. Brazil holds
the largest populations of jaguar and puma in the New
World. Cattle ranching is a major industry, and thus thisAnnual budget: V276

Parl111:

Cat ActionPlanstudied cats; and to evaluate species status in terms of POPulation distribution and protected area coverage.

country is an appropriate place to test various management
solutions to the problems of livestock predation by big catsin tropical America.
Under the auspices of the Brazilian governmental
Instituto Brasileiro de Meio Ambiente e dos Recursos
Naturais Renovaveis (IBAMA),a National Center for
Research, Management and Conservation of Predators in
Brazil has been established under the direction of an expert
in big cat ecology. The Center will coordinate and act asa catalyst for field studies, reintroduction and translocation
projects, education programs, fund raising, compensation
schemes, and country-wide data collection on carnivores.

Description:
A major conservation issue for large cat
species is that isolated reserves may not be of sufficient
size to harbor minimum viable populations. Although thetheory behind MVP is well-developed, it has not yet been
applied in a useful way to the ecology and distribution ofcats. Cat conservation could be much more strategic if(1) the MVP concept were evaluated in the specific light ofcurrent knowledge of cat biology and social organization;(2) species- and habitat- specific definitions of MVP size
were developed; and (3) the current population distribution
and protected area coverage were evaluated for eachspecies in terms of conservation of MVPs.
A team of cat specialists will conduct a workshop to
discuss the concept of MVP in relation to big cats, and to
define MVP size for the better-studied species (including
tiger, snow leopard, cheetah, lion, jaguar, leopard, andpuma). Workshop participants will also evaluate species
status based on MVP definitions by reviewing information
on population distribution and protected area coverage.

Annual budget: IV
Time frame: OngoingContact: Peter Crawshaw, IBAMA, Parque National do
Iguacu, C.P. 750, Foz do Iguacu, Parana 85851-970,
Brazil.

Research
Annual budget: V
7.

A guide to census proceduresfor cat populations
Timeframe:

Objective: To select, refine, and recommend standardized
census procedures for different cat species and habitat
types.

9.

Description: Cats are notoriously difficult to census, andestimating density, either in absolute or relative terms, isimportant for assessing the status of cat populations. The
Cat Specialist Group will work with experts in the field ofcounting animals, who will be given the opportunity tovisit various field projects and communicate with members regarding problems with censusing cat species in various habitat types. Guidelines will be produced to inform
field scientists about the strengths and weaknesses of various census techniques for cats, and to recommend which
technique is most appropriate for different cat species,
habitat types, and field conditions.

A fund for field collection and processing of
biological samples for genetic, morphological,
and clinical analysis

Objective: To establish a Cat Specialist Group Fund inorder to increase field collection and processing of biological samples for genetic, morphological, and clinicalanalysis.
Description: As described in the Research chapter, knowledge of inter- and intraspecific diversity, population
dynamics, and the ecological role of disease would begreatly enhanced by collection of more biological samples from across species ranges. Appendix 2 sets out aprotocol for field collection of such samples. Processing
and shipping can be prohibitively costly for small projects
and for individuals, and under such circumstances thesecosts will be supported by a small Cat Specialist Group
Fund.

Annual budget: IV
Timeframe: 2-3 years

8.

I yearAnnual budget: I

A workshop to define minimum
viable population size for cat species

Time frame: Ongoing
Objectives: To hold a Cat Specialist Group workshop to
define minimum viable population (MVP) size for well-277

Part III: Cat Action

PlanTime frame: Ongoing

10. Systematics of the genus FeZisand hybridization problems*

Contact: Marc Artois, CNEVA-Laboratoired’Etudes surla Rage et la Pathologie des Animaux Sauvages, BP 9,54220 Malzeville, France.

Objectives: To revise the systematics of cats of the genus
Fe&; and to investigate the present and historical incidence of hybridization between these cats and the domestic cat.

Trade
Description: Hybridization occurs widely between domestic cats and wildcats, but the extent of the problem is notknown for other close relatives included in the genus FeZis.
This museum-based study will examine the problem
through morphological, geographical, and temporal variation in the skins and skulls of Felis species. This examination will also help define subspeciation in these cats,with taxonomic evaluation of the wildcat, the widest ranging cat species, of particular importance. In addition, amodel will be developed to simulate the effects ofhybridization with domestic cats on the genetic integrity ofwild populations. This project will be complementary to
Projects 15 and 43.

12. Understanding the market
for tiger bone medicines*
Objectives: To evaluate the effectiveness of trade bans on
the markets for tiger bone medicines in China, Taiwan,
South Korea, and major overseas Asian communities; andmake further recommendations to bring illegal trade undereffective control.
Description: Commercial poaching for tiger bone is theprevailing threat to the survival of the species, yet there islittle available information on the market for tiger bone
medicines. Most major producer and consumer nations
had banned trade in tiger bone medicines by 1993, but illegal trade continues. Very little is known about consumer
demand for tiger bone, and there have been no serious
attempts to address it in terms of public education campaigns or provision of substitutes. In order to halt commercial tiger poaching effectively, conservationists must
understand the organization and dynamics of the tiger
bone market.
This project will have three components. The first is asurvey of manufactured medicines (made mainly in China
and South Korea). Samples will be collected and analyzed
by the U.S. Fish and Wildlife Service’s Wildlife Forensics
Laboratory for the presence of tiger bone. China andSouth Korea, the two major producing countries, recently
banned manufacture of these medicines; the effects of thisban will be assessed. The second component is a survey of
traditional pharmacies in major Asian and western cities
for the availability of raw tiger bone and tiger bone medicines, authentic and fake. Information will be collected
through interviewson sale volume and substitutes.
Finally, consumers of tiger bone medicines will be surveyed, so that the nature of the demand can be evaluated,
and appropriate measures formulated to address it.
Recommendations will be made for bringing the market
under effective control.

Annual budget: III
Timeframe:

3 years

Contact: Andrew Kitchener, Royal Museum of Scotland,
Chambers St., Edinburgh EHl 1JF, U.K.

11. A survey of disease in wild cat populations*Objectives: To gather and exchange published and unpublished information on the prevalence and impact of disease, particularly viruses, on cat populations, particularly
those which are small and isolated; to create a surveillance
network; and to prepare action plans with the help of computer modelling.
Description: The Laboratory for Studies of Rabies andWild Animal Pathology, sponsored in part by the Frenchgovernmei
Veterinary
review andin wild cat
offer free analysis and diagnostic services to fieldresearchers. It will aid publication of findings through
SSC newsletters and specialized meetings. A computermodel of the effects of disease in isolated populations ofcarnivores is being developed. The funds will be used tosupport graduate student thesis work and increase the center’s ability to process samples.

Annual budget: III
Timeframe: 2 years
Con tact: TRAFFIC International, 2 19c Huntingdon Road,
Cambridge, CB3 ODL, U.K.

Annual budget: I

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Part 111: Cat Action

13. The leopard cat in China: ecology andmanagement for sustainable utilization*

Specialist Group-linked fund for field projects, based on
the total number of cats in their collections. In return,annual progress reports on field projects sponsored by thefund will be provided to contributing zoos, so that appropriate educational and promotional displays can be created.

Objective: To collect and analyze the harvest, trade, andecological data necessary to establish a sustainable management system for the leopard cat in China.

c

Plan

Annual budget: I (for staff time to establish and administer the fund)Description: The leopard cat is the only species in the current international fur trade which is exported mainly froma developing country. It has been the most heavily tradedcat species over the late 1980s with the vast majority ofskins originating in southern China. The biological impactof the harvest is unknown but believed to be significant,and the Chinese government wishes to implement a more
effective management system.
The project, initiated in 1992 by the Chinese government with the support and approval of CITES, has twocomponents. The first is an intensive survey of China’s
leopard cat harvest, regulatory management, and market
organization at local, provincial and national levels. The
second component gathers ecological data on the leopard
cat in China through intensive comparative field studiesof two populations: one protected and one harvested. The
results of this study will be used to design a management
system which ensures that future leopard cat harvests are
sustainable.

Time frame: Ongoing

15. Evaluation of subspeciation and establishmentof regional captive breeding programs for the
wildcat*Objectives: To determine the degree of subspeciation in
the wildcat FeZis sihestris and to maintain viable populations of these subspecies in captivity.
Description: The wildcat has the largest range of all the cat
species, and is probably differentiated into a number ofsubspecies or perhaps even separate species. Hybridization with domestic cats is taking place across most of thisrange. It is important to conserve viable populations ofpure wildcat subspecies in captivity, while solutions to
the problems of hybridization in the wild are worked out.
Various institutions around the world maintain small wildcat populations; additional “pure” founders may need to beacquired. Captive breeding programs will be established
on a regional basis to maintain genetic diversity. Zoos will
participate in taxonomic analysis of the wildcat by providing animals and samples for study. This project will
be complementary to Projects 10 and 43.

Annual budget: III
Time frame: 4-5 years
Contact: Jinping Yu, Dept. of Zoology, 331 FunchessHall, Auburn University, Alabama 3 18495414, U.S.A.

Annual budget: IV

Cats in Captivity

Time frame: 5 years

14. Establishment of a zoo-based fund forfield conservation of wild catsContact: IUCN/SSC Conservation Breeding Specialist
Group, 1210 1 Johnny Cake Ridge Rd., Apple Valley, MN
55 124, U.S.A. AZA Felid Taxon Advisory Group, Jill
Mellen, Co-chair, Washington Park Zoo, 4001 S.W.
Canyon Rd., Portland, OR 97221-2799, U.S.A.

Objective: To establish a fund for in situ cat conservation
based on small annual donations from zoos holding catsin their collections.
Description:
Zoos are increasingly committed to andinvolved with wildlife conservation.
As discussed in
Chapter 5, zoos can make an immediate and tangible contribution to cat conservation through increased support offield projects such as those proposed in this Action Plan.
However, few zoos have the infrastructure, expertise, andbudgetary flexibility to fund international field conservation projects directly. This project proposes that zoos holding cats make a small annual contribution to a Cat

Reintroduction
16. Long-term monitoring of the reintroduced
Eurasian lynx population in Switzerland*
Objectives: To continue to monitor the process of population establishment of lynx in Switzerland.

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Pat-t Ill: Cat Action

Plan

Description:
This is the longest-runningstudy of theprocess of population estabIishment in a felid reintroduction. Its results are highly relevant to understanding thefactors (biological, ecological, demographic, cultural,political) which affect reintroduction of large felid predators. It consists of two parts: (1) monitoring of the dynamics and status of the two separate reintroduced lynx
populations (Alps and Jura Mountains), and (2) a fieldstudy of lynx ecology in the Jura Mountains. The monitoring program keeps track of the development of the total
range occupied by the reintroduced populations, lynx
killed or found dead, and damage to livestock caused bylynx. The complementary field study examines spatial andsocial structure, and recruitment and mortality rates.
Special attention is given to possible congenital problems
in the small Jura sub-population, which are suspected
because of unusually high losses of male kittens.

of extinction within the next 100 years, and even population supplementation will theoretically do little to minimize chronic inbreeding. This is a rare chance to test themodels empirically. The project, run by the University ofGeorgia, will continue to monitor population dynamics
by conducting censuses and using radio-telemetry.

Annual budget: IV

II. Species

Projects

Sub-SaharanAfricaAnnual budget: III
Time frame: Long-term ( 10+ years)
Contact: Robert Warren and Michael Conroy, Warnell
School of Forest Resources, Universityof Georgia,
Athens, GA 30602-2152, U.S.A.
See also Project 35 for the Asiatic lion.

Time frame: OngoingContact: Urs and Christine Breitenmoser and Simon Capt,
Swiss Lynx Project, Villettengassli 4, CH-3074 Muri,
Switzerland.

Black-footedcat (Fe/is nigripes):

18. Natural history of the black-footed
17. Population dynamics of a reintroduced bobcat
population in a small, isolated habitat block*Category

1

cat*

Objective: To continue the first study of the behavior andecology of the black-footed cat in order to gather basic biological information and better understand species status.

Objectives: To continue monitoring the reintroduced bobcat population on Cumberland Island, Georgia (82 kmz),
which is predicted by population viability models to havea high probability of extinction, in order to evaluate indetail the dynamics of a cat population confined to a small,isolated area of habitat.

Description: Three cats have been radio-collared on a private game reserve in the Kimberley region of South
Africa, and over 600 hours of observation data have beengathered so far. The study, sponsored in part by theMammal Research Institute of the University of Pretoria,
will continue to focus on the following problems: (1)home range size, density, and variation in seasonal activity
patterns; (2) diet; (3) breeding and denning behavior; (4)blood chemistry (in order to answer questions encounteredduring captive breeding: see Species Account); (5)intraspecific variation through the study of skins and skeletal material from various southern African collections,and (6) the gathering of biological samples from otherparts of the species range in cooperation with the projectdescribed below.

Description: Most of the world’s protected areas are lessthan 100 km2 in size. It is predicted that many parks andreserves are too small to support viable populations of cats,
but there is little empirical evidence of the processes thatdrive small populations to extinction in nature. The
Cumberland Island Bobcat Project is a unique opportunity to study the phenomenon of population viability in thefield from start to finish. The original population size,
and its genetic makeup, are known, and the island is typical of the size of most of the world’s protected areas, andpresents a worst-case scenario in terms of isolation ofmany of the world’s protected areas.
The reintroduction of bobcats to Cumberland Island
was set up by the University of Georgia as a model project to study small populations. Blood samples were collected for genetic analysis from all of the founder animals.
Population viability models have predicted a high chance

Annual budget: II
Time frame: 2-3 years
Contact: A. Sliwa, Mammal Research Institute, Universityof Pretoria, Pretoria, South Africa.

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Part III: Cat Action

19. Distributionof the black-footed.

Description: Working with recent forest cover maps, theproject will conduct presence/absence surveys in selectedparts of the African golden cat’s range to clarify population distribution, connectivity, and status. Where possible,
biological samples will be collected for genetic and morphological analysis. Data will also be collected on occurrence of sympatric cat species.

cat

Objective: To map the distribution of the black-footed cat
and gather information on species status.
Description:
The black-footed cat has an unusually
restricted range in southern Africa, and is generally considered rare. However, there are reportedly certain areas
where black-footed
cats are relatively
abundant.
Presence/absence surveys will be conducted in South
Africa, Namibia, and Botswana; and possible species
occurrence in Angola, Zimbabwe, and Mozambique will
be investigated. Survey results will aid range delineationand population status assessment. Data will also be collected on the occurrence of sympatric cat species, withemphasis on the wildcat and domestic cats, with which
black-footed cats may hybridize.

Annual budget: I
Time frame: 3 years

CheetahAfrican(Acinonyxjubatus):

Category

22. Support for the Cheetah Conservation

2(A)
Fund*

Objective: To promote cheetah conservation in Namibia
and other African range states, with concentration on cattle ranching lands outside protected areas.

Annual budget: I-II
Timeframe:Plan

2 years

goldencat (Profelis

aurata):Category

Description: The Cheetah Conservation Fund (CCF) wasestablished in Namibia in 1990. Its primary activity is
working with the cattle ranching community (through lectures, visits, and a newsletter) and the government to promote ways in which cheetah depredation of livestock canbe minimized. The CCF is also in a position to evaluate
the contribution to cheetah conservation of economic use
options developed by Namibia, chiefly trophy hunting andsale of live animals for international breeding programs(permitted under a CITES Appendix I quota system), butalso including tourism on game ranches, a growing industry in southern Africa. The CCF is conducting a radiotelemetry study to examine the ecology and socialorganization of cheetahs on farmland, has built up a substantial database of samples for genetic analysis and forsupport of the captive cheetah population, and is carryingout extensive public education programs.

1

20. Natural history of the African golden cat*Objectives: To continue the first study of the behavior andecology of the African golden cat in order to gather basic
biological information and better understand species status.
Description: Preliminary data on diet has been gatheredfrom a study of golden cat scats in Zaire’s Reserve deFaune Okapi in the Ituri Forest (J. Hart and M. Katembo in
prep.). Future work will use radio-telemetry to study
behavior, ecology, habitat use, and abundance. This project will yield the first scientifically gathered data on thenatural history of this species.

Annual budget: V
Annual budget: T-11
Time frame: Ongoing
Time frame: 2 years
Contact: Laurie Marker-Kraus and Daniel Kraus, Cheetah
Conservation Fund, P.O. Box 247, Windhoek 9000,
Namibia. U.S. address: c/o International Wilderness
Leadership (WILD) Foundation, 211 West Magnolia, Fort
Collins, CO 80521.

Contact: John Hart, Research Associate, WCS, Epulu via
Mambasa, P.O. Box 2 1285, Nairobi, Kenya. U.S. address:
Wildlife Conservation Society/NYZS, Bronx Zoo, Bronx,
NY 10460.

21. Distributionof the African

23. Factors limiting cheetah populations
outside protected areas*

golden cat

Objectives: To map the distribution of the Asiatic golden
cat and compile information on species status.

Objective: To compare cheetah reproductive and mortality
rates betw een populat ions living in (1) protected areas;
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Part III: Cat Action

Plan

Contact: Sarah Durant (Project Director), Institute ofZoology, Regent’s Park, London, NW1 4RY, U.K.
Tanzania address: Serengeti Wildlife Research Institute,c/o TANAPA, Box 3134, Arusha.

(2) commercial cattle ranching areas; (3) pastoralist grazing lands.
Description: Cheetah numbers in the Serengeti National
Park, Tanzania, appear to be limited by very high rates of
cub mortality caused by other large carnivores, chiefly
lions. Cheetahs may thus exist at higher densities outside
protected areas where other large carnivores have been
largely eliminated. Radio-telemetry studies using consistent methodology will compare the population dynamics
of cheetahs in the Serengeti to populations found in (1)commercial cattle ranches in Namibia or Zimbabwe and(2) pastoralist lands in Kenya or Tanzania. This data willlead to better understanding of the relative importance ofthe various factors limiting cheetah numbers in such areas.

25. African cheetah status survey*
Objective: To continue compilation of data on the distribution and status of cheetah populations ona regional basis.
Description: As part of a Ph.D. thesis project, status surveys (interviews)
have been carried out in Kenya,
Tanzania, and Uganda in east Africa, and in Malawi,
Zimbabwe, Botswana, South Africa, and Namibia in southem Africa over a two-year period. To complete coverageof these two regions, the project executant will conduct
interviews next in Zambia, where cheetahs are believed tobe more rare, but there is little recent available information.
Survey results will form the basis for a detailed population distribution map and prioritization of conservation
action. Data analysis will focus on determining which specific combinations of ecological and anthropogenic parameters allow healthy cheetah populations to survive.

Annual budget: III
Time frame: 4-5 years
Contact: Karen Laurenson, Upland Research Group, The
Game Conservancy, Crubenmore Lodge, Newtonmore
PH20 lBE, U.K.

24. Support for the Serengeti Cheetah Project*

Annual budget: III

Object ive: To c ontinue the long-term study of the ecol%Y of cheetahs in the Serengeti ecosystem.

Time frame: 2 years
Contact: Paule Gros and Tim Caro, Wildlife and Fisheries
Dept., University of California, Davis, CA 95616, U.S.A.

Description: In cooperation with the Serengeti ResearchInstitute, the Serengeti Cheetah Project was initiated in
the late 1970s. The project provides detailed information
on demographic rates, lifetime reproductive success andmovements of individually known cheetahs living in
Tanzania’s Serengeti National Park. It is the only suchdata set for cheetahs, and is important for purposes ofcomparison with cheetahs living in other ecosystems. The
project’s current activities are as follows: ( 1) collection ofdemographic records on known cheetahs; (2) establishment of a new study area for long-term research on woodland cheetahs; (3) assessment of relative merits forcheetahs of woodland and plains habitat; (4) determinationof the means by which cheetahs coexist with their maincompetitors, lions and hyenas; (5) using data gathered forl-4 to construct computer models for predicting minimum
reserve sizes for viable cheetah populations in different
habitats and for varying densities of other predators.

26. Distribution, status, and captive
breeding of cheetah in Zimbabwe*
Objectives: To update presence/absence surveys conducted several years ago and to capture problem animals
on commercial cattle ranches for inclusion in a captivebreeding program at Chipangali Wildlife Trust.

Annual budget: II

Description: The Chipangali Wildlife Trust is concernedthat as a result of the legal hunting of cheetahs on licensein Zimbabwe, under a quota system, and the fact that cheetahs are still killed illegally in the country, the population
could be severely reduced in the near future. This projectwill conduct presence/absence surveys and compare theresults to those carried out in the mid- 1980s and will build
up the Trust’s captive cheetah population through incorporation of wild problem animals.

Time frame: Ongoing

Annual budget: II
Timeframe: 5 years282Pat-f III: Cat Action

Plan

Contact: Vivian Wilson, Chipangali Wildlife Trust, P.O.
Box 1057, Bulawayo, Zimbabwe.

29. Predator-prey
relationship between lions
and large ungulates in South Africa’s
Kruger National Park*

27. Cheetah census and disease
surveillance in Kenya*

Objective: To measure the impact of lion predation on preyspecies.

Objectives: To conduct presence/absence surveys forcheetahs in Kenya and estimate population size; and toestablish the prevalence and importance of scabies/mange
and other diseases in wild cheetahs and methods of treatment.

Description: It is important for the management of the
Kruger ecosystem, which contains one of the largest protected lion populations in Africa, to understand the relationships between predators and prey. Lions are thedominant large predators in Kruger. A doctoral student
will conduct radio tracking and direct observations oflions, as well as aerial surveys of prey populations and distribution. The data will be used to test and refine theoretical models of lion-prey interaction (Mills and Shenk
1992).

Description: The Kenya Wildlife Service, through its field
personnel, will collect data on cheetah distribution andnumbers, as well as field samples for analysis of exposure
to disease. Treatment and evaluation of effect will be tried
out.

Annual budget: I
Annual budget: I
Time frame: 2-3 years
Time frame: l-2 years
Contact: P.J. Funston and Gus Mills, Kruger National
Park, Private Bag X402, Skukuza 1350, South Africa.

Contact: Richard Kock, Kenya Wildlife Service, P.O. Box4024 1, Nairobi, Kenya.

30. Support for the Serengeti Lion Project*
Lion (Panthemlea): Category

28. Pan-African

lion survey*2(A)Objectives: To continue the long-term study of lion behavior and ecol .ogy in the Serengeti ecosy stem.
Description: The Serengeti Lion Project is the longest-running and most extensive study of wild cats in the world.
Detailed demographic records have been compiled on
individual lions from two different Serengeti habitats: theplains and the Ngorongoro Crater. The Project has provided baseline data on many aspects of lion biology,
including the evolution of sociality, and made an important
contribution to wider issues in cat conservation, including
population genetics and the effects of disease upon population dynamics. The Project will continue to study thesesubjects through weekly monitoring of 18 Serengeti lion
prides and regular collection of biological samples.

Objectives: To continue to conduct presence/absence surveys (mail questionnaire, field surveys in key areas) toobtain a detailed distribution map of lion populations in
Africa.
Description: Lions are increasingly confined to protectedareas, and unprotected blocks of habitat which still harborlarge or significant lion populations need to be identified.
The African Carnivore Survey has compiled survey dataon lion distribution in southern Africa, and will focus next
on west African countries. East Africa will be surveyed
largely by mail questionnaire. The result will be a detailed
map of lion population distribution to aid prioritization ofconservation action.

Annual budget: V

AnnuaZ budget: III

Time frame: Ongoing

Time frame: 2 years

Contact: Craig Packer (Project Leader), Dept. of Ecology,
Evolution and Behavior, University of Minnesota, 1987
Upper Buford Circle, St. Paul, MN 55108, U.S.A.

Contact: Chris and Tilde Stuart, African Carnivore
Survey, P.O. Box 96, Nieuwoudtville 8 180, South Africa.

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Plan

31. Support for the N!aiuh,Project
the Namibian Kalahari*ture and livestock husbandry. The extent of livestock
depredation will be investigated and quantified, and various management solutions tried out. Local involvement
will be key to the success of this project, the results ofwhich will be highly relevant to the survival of the lion
outside protected areas elsewhere on the continent.

in

Objectives: To continue training Ju/hoan Bushman trackers to act as problem animal control officers in northeastern Namibia; and to develop revenue-raising options for
local people as an incentive to conserve big cats.

Annual budget: II-III
Description: As part of the Namibian government’s developing sustainable use of wildlife program for northeastern
Namibia, the project will continue to work with local
trackers to solve problems of lion predation on livestock.
Stock-raiding lions are tracked down and radio-collared,
and their movements monitored. The area is home to
Bushman communities whose traditional knowledge of
wildlife is in danger of being lost. The project aims todevelop solutions which will allow rudimentary cattle raising to develop alongside healthy populations of wild ungulates and large predators, and to work with the community
to develop economic use options based on use of wildlands and wild animals (with trophy hunting of leopards
currently being the most feasible), which will provide
badly-need revenues to the community. The project inBushmanland is an example of a growing movement in
Africa to develop economic use options for wild lands, andto cede control and management of wildlife to local residents. This project will be a test case of whether the conservation of dangerous problem animals such as lions canbe assured under such conditions.

Time frame: 3 years

Serval (Leptailurus
serwal): Category 3
No projects suggested for sub-Saharan Africa, althoughsome data on distribution and status will be collected in thecourse of surveys for sympatric cat species (Projects 19
and 21). For the north African serval, see Project 40.

Leopard(Panthera

par&m):

Category4(A)33. Evaluation of the effects of sport hunting
on leopard population dynamics
Objectives: To evaluate the impact of sport hunting on
leopard populations and test the leopard population model
developed by Martin and de Meulenaer (1988); and to
make recommendations for management of leopard sport
hunting.
Description: Leopard sport hunting is carried out on concessions in several African countries, but the effects on
population structure and dynamics have never been studied. Sport hunting is viewed as the major component of asustainable use leopard conservation strategy on the subcontinent. Leopard population dynamics and response tovarying degrees of offtake have been modeled by Martinand de Meulenaer (1988), but their models have not yet
been field-tested. This radio-telemetry study will compare
the population dynamics of a protected population of leopards to one that is subject to sport hunting. The project will
also test the hypothesis that leopards exhibit complete
compensation.

Annual budget: III
Time frame: 3 years
Contact: Philip Stander, Ministry of Wildlife, Conservation and Tourism, P.O. Box 17, Grootfontein, Namibia.

32. Resolving lion and livestockconflicts in west Africa
Objectives: To evaluate the extent of livestock loss to lions
in a west African country; to evaluate the threat posed tolion populations by persecution; and to develop appropriate management solutions applicable to the region.

Annual budget: III
Time frame: 2-3 years

Description: Lions are patchily distributed through thewooded savannahs of west Africa. This region is an
important part of the species range: while the lion has not
yet been eliminated from private lands and restricted toprotected areas, this process is well under way. The project will select a west African study area which has a relatively large population of lions occurring on private lands
where the predominant form of land use is mixed agricul-

34. Leopard ecology and density intropical African rain forest*
Objectives: To continue study of the natural history of
leopards in African tropical rain forest, and estimate their
numbers in this habitat type.
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Description: Tropical rain forest makes up a large part ofthe leopard’s sub-Saharan range, but little is known of theirnatural history in this type of habitat. It has been predictedthat leopards are most abundant in tropical rain forest, if
density increases with rainfall (Martin and de Meulenaer
1988). Continuing study of leopards in the Ivory Coast’s
Tai’ National Park and the Ituri Forest in Zaire will provide
important data for understanding the status and ecology ofthe leopard. In addition, blood samples will be collected foranalysis of genetic differentiation and diversity.

Planability of wild Asiatic lions and (2) relieve the problem oflion depredation in the Gir Forest. A recent PHVA workshop (Walker 1994) evaluated potential release sites andselected Palpur Kuno Wildlife Sanctuary as the mostpromising. In cooperation with the Wildlife Institute ofIndia and the state and national governments, a feasibility
study is being carried out. Lions moved from the Gir to therelease site would be genetically sampled and marked orradio-collared.
Annual budget: V

Annual budget: II
Time frame: 3-5 years
Time frame: 2-3 years
Contact: Ravi Chellam and A.J.T. Johnsingh, Wildlife
Institute of India, P.O. Box 18, Chandrabani, Dehra Dun
249 001, Uttar Pradesh, India.

Contact: Ivory Coast: Frederic Dind, B.P. 20, Part
National de Ta& Cote d’Ivoire. Switzerland address: c/o
Christophe Boesch, Institute of Zoology, University ofBasel, Rheinsprung 9, Basel. Ituri forest, Zaire: John Hart,
Research Associate, WCS, Epulu via Mambasa, P.O. Box
21285, Nairobi, Kenya. U.S. address: Wildlife Conservation SocietylNYZS, Bronx Zoo, Bronx, NY 10460.

Cheetah

36. Conservation

lion (I? lea per&@:

Asia

Category1 (A)of the Asiatic cheetah in IranDescription: The Iranian population is the only remaining
cheetah population in Asia and the largest in the northAfrican-southwest
Asian region, but it is in serious danger of extirpation, with numbers estimated at fewer than
50. These cheetahs may be different-at
least in terms ofnatural history, and in terms of morphology and geneticsfrom sub-Saharan cheetahs. The population is fragmented,
low density, and declining from habitat degradation andpersecution of both cheetahs and their antelope prey. In
the first year, this project will map the distribution of cheetahs in Iran; collect samples for molecular analysis; prioritize populations for conservation;and produce a
conservation management plan. In the second year andthereafter, recommendations for action will be carried out.

African wildcat (Fe/is silvestris,
lybica group):
Category 5
No specific projects suggested for this region, but see
related Projects 10, 15, and 43. Some distribution data will
be gathered by Project 19.

Asiatic

Category

Objectives: To develop and implement a conservation
management plan for the cheetah in Iran.

Caracal (Camcal caram/):Category 4
No projects suggested for this region, but see related
Project 42 in India.

North Africa and Southwest(A. juhtus):

1

35. Establishment of a second
population of Asiatic lions in India*

Annual budget: I-IV
Time frame: 3-5 years

Objective: To establish a second population of Asiatic
lions in a selected reserve in India.

37. Natural history, genetics, and conservation
status of the cheetah in the Saharan region

Description: India’s Gir Forest holds the world’s only population of Asiatic lions. The lions appear to be at maximum density, and there are continual problems with lions
emigrating into settled areas, killing both livestock andpeople. The single population, while large, is still vulnerable to stochastic events, and establishment of a second
population should both (1) substantially lessen the vulner-

Objectives: To conduct the first study of the behavior andecology of the cheetah in the Saharan desert region; tocollect samples for genetic analysis; and to assess its conservation status.

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Description: The cheetah is the last big cat to survive in the
Saharan desert, where the habitat is very different from theplaces cheetah ecology has been studied, in east and southern Africa. Cheetahs are very sparsely distributed in the
Sahara, with populations centered around mountain
ranges; there may be fewer than 200 animals across theentire desert region. This project will examine the physiological and behavioral adaptations which have enabledcheetahs to persist in the Sahara, evaluate intraspecificdiversity through genetic comparison with east and southem African animals, and aid assessment of the cheetah’sconservation status throughout the region by providing
baseline data on ecology, prey requirements, abundance,and such human-related factors as livestock depredation.
Appropriate study areas include the Tassili and Hoggar
mountains of southeastern Algeria, or the Termit Massif
region in northeastern Niger.

Leopard(I? pardus):Category3(A)39. Distribution and conservation status of
the leopard in north Africa and southwest

Asia*

Objectives: To hold a workshop of regional cat specialists
to map out current distribution of the leopard in the region,assess population status, and coordinate conservation
action.
Description: Leopards are the last big cats to survive inany numbers in the region, but their distribution is highly
localized, with small populations in low numbers. It isnecessary to map out population distribution and prioritize conservation action, especially lobbying for creationof reserves in key areas. A workshop will be held inviting professionals from north African and southwest Asian
countries, including Turkey and the Caucasus republics.
The workshop will issue recommendations for conservation action by national authorities, and will establish aregional network of professionals to improve coordination and information sharing.

Annual budget: II
Time frame: 2-4 years
See also related Project 78.

Annual budget: III
Serval (L. serval):

Category2(A)Time frame: 1 year

38. Survey for the leopard and the servalin the Atlas Mountains of Morocco*

Contact: Peter Jackson, Cat Specialist Group Chairman,
1172 Bougy, Switzerland.

Objective: To identify whether leopards and servals still
exist in the Atlas Mountains of Morocco and selected partsof northern Algeria.

40. Support for the Arabian Leopard Trust*Objectives: To promote conservation of the leopard (andother felids) on the Arabian peninsula through governmental lobbying, public education, and captive breeding.

Description: Leopards and servals are rare and possibly
extinct in the humid scrub and mixed woodlands of northAfrica. These animals are likely to have been isolated
from populations south of the Sahara for at least 7,000
years and perhaps much longer, and represent an important
potential reservoir of intraspecific diversity. In cooperation with the governments of Morocco and Algeria and the
Rabat Zoo, presence/absence surveys will be conductedusing interviews as well as confirmatory techniques.

Description:
Based in the United Arab Emirates, theArabian Leopard Trust was set up in 1993. Its activities
focus on: (1) public education campaigns about the threatened status of the cats of the Arabian peninsula, particularly the leopard; (2) improving regional management of
captive cat populations; (3) lobbying for legal prohibitionof cat hunting; (4) lobbying for reserve creation; (5) assessing the feasibility of reintroducing large antelope preyspecies raised in captivity (as was done successfully for the
Arabian oryx).

Annual budget: II
Time frame: 1 year
Contact: Haddane Brahim, Part Zoologique National de
Rabat, B.P. 4142, 12000 Temara, Morocco. Switzerland
address: Cat Specialist Group Chairman, 1172 Bougy,
Switzerland.

Annual budget: III
Time frame: OngoingContact: Marijcke Jongbloed, Arabian Leopard Trust,
P.O. Box 12119, Dubai, United Arab Emirates.

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Sand cat (Fe/is margarita):
41. Distribution

Category

Contact: S.P. Goyal, Shomita Mukherjee, and A.J.T.
Johnsingh, Wildlife Institute of India, P.O. Box 18,
Chandrabani, Dehra Dun 248 001, Uttar Pradesh, India.

4of the sand cat

Objectives: To map the distribution of sand cat populations
and assess species status in key parts of its range.

Jungle cat (Fe/is chaus): Category 5a
No projects suggested for this region. Some information
on distribution will be collected by Project 80.

Description: It is not clear whether major gaps in the sandcat’s range occur in northeastern Africa and Iran, and thestatus of the southeasternmost population in the deserts of
Afghanistan and Pakistan is unknown. Presence/absence
surveys in selected areas of western Egypt, Libya, Chad,and Sudan will be overseen by a Cat Specialist Groupregional coordinator. Surveys will also be conducted inselected parts of Iran, the Registan Desert of Afghanistan,and selected parts of Baluchistan, Pakistan (see Project 78).

WildcatCaracalCategory

lybica group):

Category

5b

Objectives: To map the distribution of wildcat populations in Israel; to assess the degree of hybridization withdomestic cats; to study the behavior and ecology of “pure”
wildcats in the wild; and to improve the captive breedingof “pure” wildcats at the Tel Aviv University Researchzoo.

2 years(C. caracal):(E. silvestris,

43. Natural history, status and captive
breeding of the wildcat in Israel*

Annual budget: II
Timeframe:

Plan

Description: Wildcats are becoming increasingly rare insouthwest Asia, and the degree of hybridization with thedomestic cat has not been well assessed for this region.
This is of special interest, as it is in this region that the
wildcat was probably originally domesticated. The behavior and ecology of “pure” wildcats has not been studied
across Zybica’s entire range. Tel Aviv University maintains an important captive population of wildcats,
descended from founders captured nearly 40 years ago.
Their facilities need to be improved, and the offspring circulated in an international captive breeding program (seerelated Project 15).

5a (A)

42. Natural history, distribution, andstatus of the caracal in India*
Objectives: To map caracal presence/absence, conduct a
detailed field study of its behavior and ecology, and assess
species status at the eastern limits of its range.
Description: The caracal is believed to be rare in India, andin all of its Asian range natural history has been studied
only in an agricultural area of Israel. In cooperation with
the Wildlife Institute of India and State Forestry
Departments, the project executant will conduct presence/absence surveys for the caracal in India, including
both protected areas and key areas of unprotected habitat,
notable among them Kutch division of Gujarat, theGwalior division of Madhya Pradesh and the Aravalli Hillsof Rajasthan. Distribution will also be collected for othersympatric small cat species. Survey results will form thebasis for population distribution maps, and will aid better
understanding of species status. Meanwhile, the ecology
and behavior of the caracal in this region will be the focus
of a radio-telemetry study in the Sariska Tiger Reserve,
Rajasthan.

Annual budget: III
Timeframe: 5 years
Contact: Heinrich Mendelssohn, University of Tel Aviv,
Faculty of Life Sciences, P.O. Box 39040, Ramat Aviv
69978, Tel Aviv, Israel.

TropicalAsia

Tiger (Panthera

Annual budget: II

Tigris): Category

1 (A)44. Protection of the Amur tiger and its
habitat in the Russian Far East*

Time frame: 2-3 years

Objectives: (1) To support anti-poaching operations toimprove protection of the largest remaining population of287

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Description: The first study of the Amur tiger, using snow
tracking and radio-telemetry, began in 1991 in Russia’s
Sikhote Alin Biosphere Reserve. Russian and American
researchers from the U.S.-based Hornocker Wildlife
Research Institute are collecting baseline data which will
be used to document the tiger’s spatial organization, habitat utilization, and population dynamics. Understanding ofthese variables will facilitate the creation of an appropriate reserve system for the Amur tiger, which occurs mainly
outside of protected areas. This project works in close
cooperation with Projects 44 and 67.

Amur tigers; (2) to conserve,a sufficient amount and arrayof high quality habitat for tigers and other wildlife to maintain viable populations and ecosystem functions over the
long-term; and (3) to increase conservation and environmental awareness throughout the region through public
education initiatives.
Description: Amur tigers in Russia, home to most remaining animals of this subspecies, have come under intensive
poaching pressure in the last few years. There are perhaps no more than 150-200 remaining. This project has
been put together by a broad coalition of NGOs to support the Russian government in its efforts to protect thetiger. Activities include the following: ( 1) strengthening
the ranger forces of the Lazovskiy and Sikhote-Alin
Nature Reserves; (2) creating and supporting mobile federal anti-poaching brigades on a regional level and local
community game guard patrols to safeguard tigers outside protected areas; (3) assisting Russian Customs withtechnical training to improve enforcement of trade bans;(4) publicizing the strengthening of anti-poaching measures; (5) improving protected area coverage for the tiger
by expanding existing areas, establishing new reserves,
creating conservation management areas and protecting
corridors between them; (6) monitoring the impact of timber and mining extraction on tiger populations andpromoting sustainable forestry practices; (7) using Geographic Information System (GIS) map databases to assist
these processes; (8) working with Russian NGO counterparts to broaden public awareness and support for tiger
conservation. Cooperating organizations include: Ministryof Environmental Protection and Natural Resources of theRussian Federation, the Pacific Institute of Geography andSoils, WWF-Germany, WWF-U.S., the Primorski Regional Association of Indigenous People, the Hornocker
Wildlife Research Institute (U.S.), the Tiger Protection
Society (Russia), the Tiger Trust (U.K.). and the Siberian
Tiger Conservation Network (U.S. ).

Annual budget: V
Time frame: 2-3 years
Contact: Dale Miquelle, Sikhote-Alin Reserve, Terney,
Primorye Territory, Russia.
U.S. address: Howard
Quigley, President, Hornocker Wildlife Research Institute,
P.O. Box 3246, University Station, Moscow, ID 83843.

46. Conservation

Objectives: To develop and implem .ent a conservation
management plan for the South China tiger.
Description: The South China tiger is not only the mostprimitive tiger subspecies, and probably ancestral, but also
the rarest, with only 30-80 animals left in highly-fragmented habitat in south-central China. It will almost certainly disappear in the very near future unless active effort
is made to conserve it, as the present sub-populations areprobably too small, fragmented, and threatened to persist
in the long-term. This project, sponsored in part by the
Chinese government, will develop and implement a conservation management plan for the subspecies. Activities
will include support for improved reserve management in
key protected areas, creation of new reserves to preserve
habitat corridors between sub-populations, and a variety ofsocio-economic development projects to reduce conflict
between local people and wildlife authorities.

Annuul budget: V
Time frame: 5 years
Contact: Georg Schwede, WWF Germany.
701127.6000 Frankfurt A/M 70, Germany.

of the South China tiger*

Annual budget: V

P.O. BoxTime frame: S- 10 years
Contact: Gui Xiaojie, Hunan Provincial Dept. of Forestry,
39 Nan Da Road, Changsha 410 007, Hunan, China.
Meng Sha, Ministry of Forestry, Hepingli, Beijing 100
7 14, China. Switzerland address: Cat Specialist Group
Chairman.

45. Behavior and ecology of the Amur tiger*
Objectives: To continue the first radio-telemetry study of
the Amur tiger in Russia’s Sikhote-Alin
Biosphere
Reserve.

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47. Support for Global Tiger Patrol*

Time frame: 1 year

Objectives: To improve conservation of the tiger throughout its range by building support among people living outside tiger reserves through community development
initiatives, and by establishing an award/incentive programfor anti-poaching personnel.

Contact: Valmik Thapar, Cat Specialist Group ViceChairman for Asia, 19 Kautilya Marg, New Delhi 110 02 1,
India.

49. Distribution, abundance, and ecologicaland conservation status of the tiger in India*

Description: The tiger is seriously threatened by commercial poaching pressure throughout its range. India holds
the largest population of tigers and has invested heavily intheir conservation. A massive effort is needed to protect
tiger populations from catastrophic losses to poaching.
The Global Tiger Patrol, formerly the Ranthambhore
Society, was established in the U.K. in 1992 to set up community development programs around key protected tiger
populations to encourage local support for their conservation and thereby reduce poaching. Global Tiger Patrol isalso setting up, with the cooperation of the Indian government, an award scheme for game guards who capture
poachers. This project, partly supported by the European
Union, is actively expanding Global Tiger Patrol’s activities to other tiger range states

Objective: To evaluate the status of the tiger in Project
Tiger reserves and other areas.
Description: Despite a series of all-India censuses from1972- 1993, there is still considerable uncertainty about theexact distribution and number of tigers in India, most ofwhich live outside the 21 Project Tiger reserves. This
information is essential in order to assess the survival
prospects of the tiger in India and to improve conservation management. Basic ecological data on habitat andprey availability outside the special reserves is lacking.
Rigorous techniques for assessing these parameters havenot been developed and applied. These lacunae were highlighted by the government of India during the international
symposium on tiger in New Delhi in 1993. In northern
India, the project executant will visit reserves and othertiger areas to collect information on the distribution of tiger
populations, population structure, and reproductive andsurvival rates. In southern India, the project executant willconduct 16 weeks worth of field work at each site, with theaim of developing a standardized methodology for measuring habitat quality, tiger numbers and prey densities,
as well as predictive models to evaluate tiger population
status based on these parameters. There will be close collaboration with Project 48.

Annual budget: V
Time frame: OngoingContact.- Peter Lawton, Director, Global Tiger Patrol, 19
Kautilya Marg, New Delhi 110 02 1, India.

48. Improving the reliability
tiger censuses in India*

of

Objective: In collaboration with the Government’s Project
Tiger Directorate, to conduct a workshop to improve tiger
census techniques by evaluating the appropriateness fordifferent habitat types of various methods, including phototraps, statistical analysis of pugmark measurements, andtrend estimates by field staff

Annual budget: III
Timeframe: 3 years
Contact: Northern India: S. Deb Roy IFS (Retd), 24/4 Type5, Lodi Complex, New Delhi 110 003, India. Southern
India: K. Ullas Karanth, Center for Ecological Studies, 499
Chitrabhanu Road, Kuvempu Nagar, Mysore 570 023,
Karnataka, India. U.S. address: c/o NYZS/ Wildlife
Conservation Society, Bronx Zoo, Bronx, NY 10460.

Description: The Indian government has estimated tiger
numbers for the past 20 years from pugmark censuses, but
the accuracy of the technique has been challenged. The
present tiger poaching crisis demands that reliable, rapid,
and inexpensive methods of estimating tiger numbers be
developed so that the status of tiger populations can be
accurately monitored. Refined, computerized analysis of
pugmarks, photo-trap transect surveys, and trend estimatesby field staff require evaluation. The workshop will
review and field test these techniques.

50. Conservation of viable tiger populationsin India: the effects of population isolation andmovement corridors
Objectives: To evaluate, as part of Project Tiger, the viability of small and isolated tiger populations, as well as

Annual budget: III-IV

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Description: The Ranthambhore Tiger Reserve in northern
India became world famous in the 1980s for the opportunities it provided to view tigers in the wild. However, local
people have suffered from near exhaustion of natural
resources outside the reserve, as well as suffering depredation of crops by wild animals. Conflict with reserve
staff has occurred when people sought to graze their cattle and collect wood and fodder within the protected area.
Ranthambhore exemplifies the situation of many reserveswith tiger populations in the Indian subcontinent and elsewhere. The Ranthambhore Foundation was established in1988 to remedy the situation by establishing a sustainable
development program to improve the livelihood of people
living around the reserve. The Foundation works in sevenvillages, and its activities include: growing trees for firewood, timber, and fodder for livestock; rehabilitating
denuded grazing lands; providing new high-yielding cattle breeds adapted to stall feeding; organizing marketing of
milk; providing medical and family planning facilities andhealth education; organizing income producing activities
for women (e.g., traditional needlework, which is marketed in the cities); and providing environmental education
for all, but especially the children, with audio-visuals andvisits to the reserve. The success of the project has
attracted international attention and support.

the potential of corridors between them to improve viability; to map the presence of potential corridors throughout
the country and to survey these areas to confirm use bytigers; to undertake measures to ensure conservation ofimportant corridors.
Description: This is a pilot project relevant to big cat conservation. India holds the world’s largest population oftigers, and has been outstanding in its commitment to tiger
conservation. Project Tiger is based upon conservationof tigers in protected areas which are representative of thediversity of tiger habitat. Many of the reserves, as well asother areas with tigers, are isolated, and may be too small
to harbor tiger populations of a sufficient size to ensure
long-term viability. Conservation biology theory predicts
that even a very low level of immigration can substantially
improve the viability of otherwise isolated populations.
This long-term study, in cooperation with the Wildlife
Institute of India, will investigate the effects of population isolation and corridors on tiger populations and theirconservation.
The project has three components. The first is a longterm study to compare the health and dynamics of three
populations: a small isolated population; a large isolated
population; and a population undergoing occasional immigration of new animals through a corridor. Study sites will
be selected which are of similar size and habitat composition. The necessary legal measures should be taken toassure the integrity of the habitat corridor for the durationof the study, but this should not necessarily preclude
human activity. The tigers will be sampled for molecularanalysis of genetic diversity and kinship, and for reproductive parameters. The second and parallel component
will be to identify potential corridors linking key tiger populations elsewhere in India, and conduct sign surveys tovalidate actual use. The third component will be for the
Indian government to undertake any legal measures necessary to preserve the viability of corridors.

AnnuaZ budget: III
Time frame: OngoingContact: Valmik Thapar, Honorary Director, Ranthambhore Foundation, 19 Kautilya Marg, New Delhi 110 02 1,
India.

52. Support for the Tigerhaven Wildlife Trust*Objectives: To mitigate conflict between tigers of India’s
Dudhwa National Park and local people living on the park
boundaries by paying livestock loss compensation.

Annual budget: I-V

Description: Dudhwa National Park is home to an important tiger population in northern India, and there is a highlevel of conflict between people and tigers in the vicinity.
Sugarcane is grown right up to park borders, providing
attractive habitat for park wildlife, including tigers, which
often breed there. There have been numerous incidents ofhuman deaths and livestock predation, while there has been
evidence of poaching, for which there is a strong incentive
because of the current high value of tiger products for
smuggling to China. Most of India’s tiger reserves are situated in areas with high human populations, and the government’s program of compensation for loss of livestock
has not functioned smoothly. This pilot project will test theability of an NGO to deal effectively with the problem.

Timeframe: Long-term ( 10+ years)

51. Support for the Ranthambhore
Foundation’s eco-development

project*

Objectives: To support the Ranthambhore Foundation’s
exemplary project aimed at improving the living standards of local people around the Ranthambhore Tiger
Reserve, promoting their understanding of the value of
wildlife reserves, and thereby gaining their support forconservation.

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Street, P. 215 Khu Thanh Cong, Hanoi, Vietnam. Others
and Overall: Alan Rabinowitz, Asia Director, NYZS/
Wildlife Conservation Society, Bronx Zoo, Bronx, NY
10460, U.S.A. Peter Jackson, Chairman, IUCN/SSC Cat
Specialist Group, 1172 Bougy, Switzerland.

The Tigerhaven Wildlife Trust will liaise with communities on the periphery of Dudhwa National Park toestablish a pilot compensation scheme for loss of livestock,
paid from a trust fund managed by the NGO.
Annual budget: I-II
Timeframe: Ongoing

54. Support for the Chitwan

Tiger Project*Contact: Arjan Singh, Tigerhaven Wildlife Trust, P.O.
Pallia, Box 8, Dist. Kheri 262 902, Uttar Pradesh, India.

Objectives: To continue the long-term monitoring of thetiger population in Nepal’s Chitwan National Park.
Description: The project, run in cooperation with the U.S.based Smithsonian Institution and the government of
Nepal, employs two professional tiger trackers to compile
daily records on known tigers. It has provided the only
comprehensive set of long-term data on tiger population
dynamics.

53. Current distribution of tiger populations
in east and southeast Asia*
Objectives: To identify current population distribution andstatus of tigers in Bangladesh, Cambodia, Laos, Malaysia,
Myanmar, North Korea, Sumatra, Vietnam, and southwest
and northeast China.

Annual budget: I

Description: In cooperation with national governments,
Cat Specialist Group and NYZS/Wildlife
Conservation
Society coordinators will oversee presence/absence surveys for the tiger in southeast Asian range states. The status of the tiger is not well understood in these countries,
but there are fears that poaching pressure has led to widespread declines. Preliminary surveys have been carriedout in Laos, and surveys are in the process of being set up
in southwestern China, North Korea, Sumatra, Vietnam,
and Bangladesh. Similar work will also be done in otherrange states. Survey techniques will rely on networking
between regional coordinators and local wildlife officials
and other experts. Survey data will be used to compile
accurate population distribution maps and prioritize populations for conservation action. These surveys will also collect distribution information on other sympatric cat species.

Time frame: OngoingContact: Charles McDougal, Smithsonian Research
Associate, Tiger Tops, P.O. Box 242, Kathmandu, Nepal.

55. Development of a tiger
population simulation model*
Objectives: To: (1) construct a realistic computer simulation model for tiger populations based on the extensive
data set from Royal Chitwan National Park, Nepal; (2)use the model to design an efficient approach for fieldresearch; (3) use the model to explore the consequencesof poaching on long-term population viability; (4) use themodel to explore the potential consequences of inbreeding depression on long-term population viability.

Annual budget: I-II/per country
Description: The tiger population simulation model may
be used to address the following questions: How are viable
populations of tigers protected given limited knowledge?
How does one decide whether a population is thriving or in
danger of extinction ? How important is inbreeding depression from a managerial perspective? Is a given protectedarea large enough to support a viable population? Would
additions or corridors safeguard the population? When is
active intervention necessary and what are the likelihoodsof success of various types of intervention? Should limited
resources be expended on small isolated populations
which appear to be doomed to extinction? How should aresearch program be designed to study tigers to make themost efficient use of limited resources? Are there criticalparameters to monitor which would indicate the status ofa particular population?

Time frame: I-2 years
Contact: Bangladesh: Anisuzzaman Khan, Nature Conservation Movement, 13 Monipuripara Serebangla Nagar,
Dhaka, Bangladesh. Southern and Southwestern China:
Lu Houji, Dept. of Biology, East China Normal University,
Shanghai 200 062, China. Indonesia (Sumatra): Ronald
Tilson, SSP Tiger Coordinator and Director of Conservation, Minnesota Zoological Garden, 13000 Zoo Boulevard,
Apple Valley, MN 55124, U.S.A. Laos: Klaus Berkmiiller, IUCN-Laos, 15 Fa Ngum Road, Vientiane, Laos.
North Korea: Pak U 11, Director, Research Center for
Nature Protection, Academy of Sciences, Pyongyang,
Peoples Democratic Republic of Korea. Vietnam: Dao
Van Tien, Laboratory of Zoology, 19 Le Thanh Tong
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Plan

The model has been developed at the University of
Minnesota. It is individually-based,
stochastic and spatial, and includes detailed behavioral characteristics, suchas infanticide and dispersal patterns. The project is to becontinued as a graduate thesis.

study site. The project coordinator will oversee presence/absence surveys in the Bomean highlands and otherparts of the island.

Annual budget: I

Time frame: 2-3 years

Time frame: 1 year

Contact: Mel Sunquist, 118 Newins-Ziegler
Museum of Natural History, University
Gainesville, 32611 FL, U.S.A.

Annual budget: I-II

Contact: J.L. David Smith, Anthony M. Starfield, andJohn Kenney, Conservation Biology Program, Universityof Minnesota, St. Paul, MN 55 108, U.S.A.

CloudedCategory
56. Genetic evaluation of subspeciation

leopard2(A)(NeofelisHall, Florida
of Florida,

nebulosa):

in the tiger*
58. Natural history of the clouded leopard*

Objective: To evaluate subspeciation in the tiger throughrepresentative geneti c analysis.

Objectives: To conduct the first study of clouded leopard
behavior and ecology through radio-telemetry.

Description: Several molecular methodologies are being
used to re-examine tiger subspecies in terms of genetics.
Tiger subspecies have been well-studied morphologically,
but a genetic component is also necessary.
Previous
efforts have been compromised by a limited selection of
captive animals. This study is collecting biological samples from wild-caught animals in Russia, India, Sumatra,and other Asian range states, and will collect biologicalsamples from certified pure-bred South China tigers from
Chinese zoos.

Annual budget: II

Annual budget: III

Time frame: 2-3 years

Timeframe:

Contact: A.J.T. Johnsingh, Wildlife Institute of India, P.O.
Box 18, Chandrabani, Dehra Dun 248 001, Uttar Pradesh,
India.

Description: A preliminary survey by the Wildlife Instituteof India has identified the Dampa Wildlife Sanctuary ofMizoram, India, as a suitable study area. The behavior andecology of the clouded leopard have never been studied, amajor gap in natural history knowledge among the larger
cats. This study will provide the first quantitative data tohelp evaluation of species status.

2 years

Contact: Stephen J. O’Brien, Laboratory of Viral Carcinogenesis, National Cancer Research Institute, Bldg. 560
Room 21-105, Frederick, MD 21702-1201, U.S.A.

59. Distribution of the clouded leopard
and other sympatric small cats*
Borneanbay cat (Catopuma

badia): Category

1
Objectives: To map the distribution of the clouded leopard and other small cats.

57. Natural history, distribution, andstatus of the Bornean bay cat*

Description:
Regional coordinators will oversee presence/absence surveys by networking with local wildlife
officials and other experts, and the results will form thebasis of a detailed population distribution map. The component in southern and southwestern China is of particular importance due to the wide but patchy distribution of
the clouded leopard and Asiatic golden cat in that country. These surveys will also collect distribution data onother sympatric cat species, and will aid evaluation of theirstatus.

Objectives: To conduct the first study of the behavior andecology of the Bomean bay cat using radio-telemetry; andto compile information on its distribution and status.
Description: The Bornean bay cat is the least known cat
in the world. Nothing is known of its natural history. It
has a limited distribution on the island of Borneo and isapparently quite rare. Animals will be captured, radio-collared and sampled for genetic analysis in an appropriate

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Plan

Annual budget: I-II

62. Distribution

Time frame: 3 years

Objectives: To map the distribution of the flat-headed cat
and assess the species status.

Contact: Peter Jackson, Chairman, Cat Specialist Group,1172 Bougy, Switzerland; China: Lu Houji, Dept. ofBiology, East China Normal University, Shanghai 200
062, China.

Asiatic golden
Category 2

cat (Catopumaof the flat-headed cat

Description: A coordinator will oversee presence/absence
surveys for the flat-headed cat in Burma, Thailand,
Malaysia and Indonesia. Priority goes to the Isthmus ofKra (Burma and Thailand), a relatively densely settled
region which forms the northern limit of its range. Data
will also be collected on the occurrence of other sympatriccat species. Survey results will form the basis for a
detailed population distribution map.

temmincki):

60. Natural history of the Asiatic golden cat
Annual budget: I-II
Objectives: To gather the first information on the behavior and ecology of the Asiatic golden cat through radiotelemetry.

Timeframe: 2-3 years

Rusty-spotted
Category 2

Description: The natural history of the Asiatic golden cat
has never been studied. It is very difficult to evaluate thestatus of this vulnerable species lacking such information.
A small number of animals will be captured, radio-collared, and monitored in a selected study area.

cat (Prionailurusrubiginosus):

63. Natural history and distributionof the rusty-spotted cat

Annual budget: I-II

Objectives: To gather the first information on the behavior and ecology of the rusty-spotted
cat throughradio-telemetry;
and to map distribution through presence/absence surveys.

Time frame: 2-3 years
For distribution surveys, see Project 59.

Flat-headedcat (Prionaikms
Category 2

Description: The rusty-spotted cat has a limited distribution in India, but apparently occurs in a variety of habitats. Its behavior and ecology have never been studied,and its status is unclear. A radio-telemetry study will be
carried out to collect baseline data on its natural history
and habitat requirements, and presence/absence surveys
will be conducted through networking with local officials
and other experts.

planiceps):

61. Natural history of the flat-headed cat
Objectives: To gather the first information on the behavior and ecology of the flat-headed cat through radiotelemetry.

Annual budget: I-II

Description: The flat-headed cat has unusual physical features which point to a semi-aquatic existence, but it has
never been studied in the wild. A radio-telemetry study
will be conducted which looks at the cat’s ecology andbehavior in both protected wetland habitat and alteredhabitat, such as oil palm plantations. The results will
greatly aid understanding of species status.

Time frame: 2-3 years

Fishingcat (Prionailurus

viverrinus):

Category2

64. Natural history of the fishing cat*Objectives: To continue the first comparative radiotelemetry study of the behavior and ecology of the fishing
cat and other sympatric cat species.

Annual budget: I-II
Time frame: 2 years

Description: This project will yield the first scientificallygathered data on the nat ural history of the fishing cat.

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Part 111: Cat ActionPlanacters, have argued that they are not closely related. This
study will greatly aid understanding of the species.
Animals will be captured, radio-collared, and sampled forgenetic analysis at an appropriate study site.

Fishing cats will be captured and radio-collaredin
Thailand’s Huai Kha Khaeng Wildlife Sanctuary, alongwith other sympatric cat species (leopard cats and leopards
have been collared so far). This study will focus on thefood habits, habitat use and ecological coexistence of cat
species in Asian tropical monsoon forest. Data on the fishing cat will be compared to results from a study of fishingcats in Nepal’s Chitwan National Park.

Annual budget: I-III
Time frame: 3-4 years
For distribution surveys, see Project 59.

Annual budget: II
Time frame: 2-3 years

LeopardContact: J.L. David Smith, Dept. of Fisheries andWildlife, 200 Hodsun Hall, University of Minnesota, St.
Paul, MN 55108, U.S.A.

(I? pardus):Category3(A)67. Support for the Far East Leopard Fund*
Objectives: To ensure the conservation of a viable population of Amur leopards in the Russian Far East.

65. Distributionof the fishing cat
Description: The Amur leopard is the rarest form of leopard, distinctively unique with its large, open rosettes, andwith numbers estimated at only 30 animals in Russia, withperhaps a few in North Korea and China. A RussianAmerican scientific team associated with the Fund (seeProject 45) is carrying out the first radio-telemetry study in
the Kedrovaya Pad Nature Reserve in Ussuri province.
Funds raised by the Far East Leopard Fund will be used
to increase anti-poaching protection in the reserve, to initiate a captive breeding program, and lobby for the creationof a new reserve in important leopard habitat.

Objectives: To map the distribution of the fishing cat andassess the status of populations in key large wetlands.
Description: The fishing cat is strongly associated withwetland habitats, and is expected to have a patchy distribution over its range. It may possibly have become extinct
along the coast of southwestern India; this area is of toppriority for surveys. A Cat Specialist Group regional
coordinator will oversee presence/absence surveys in thisand other key areas of the fishing cat’s range. This will
involve networking with local wildlife officials and otherexperts, as well as site surveys in selected areas. Survey
data will form the basis of a detailed population distribution map. Data on the occurrence of sympatric cats will
also be collected.

Annual budget: I-III
Time frame: OngoingContact: Viktor Korkishko,
Far East Leopard Fund,
Kedrovaya Pad Reserve, Primorsky Krai 692 710, Russia.
Switzerland address: Cat Specialist Group Chairman.

Annual budget: I-II
Time .frame: 2 years

68. Leopard ecology in three habitats in India*
Marbledcat (Pardofelis

marmorafa):

Category

2
Objectives: To gather basic information on the natural history of leopards in three different Indian habitats.

66. Natural history of the marbled cat

Description: The leopard is the most common large cat outof the five found in India, but next to nothing is known of
its behavior and ecology. This information would be valuable for improving management of leopard populations.
Preliminary predation data has been gathered from the Gir
forest in Gujarat; the two other study sites include
Mundanthurai plateau in Kalakad-Mundanthurai
Tiger
Reserve, and Garhwal-Kumaon
Himalaya in the state of
Uttar Pradesh, where numerous leopards are killed every
year as a result of conflict with humans.

Objectives: To gather the first information on the behavior and ecology of the marbled cat using radio-telemetry.
Description: The marbled cat is among the most mysterious in the family Felidae. Nothing is known of its natural
history or habitat requirements; capture of animals forstudy is impeded by their apparent strong arboreality. Its
fur pattern is remarkably similar to the sympatric clouded
leopard, although several authorities, based on skull char-294

Part 111: Cat Action

Contact: Ryuchi Masuda, Naoyi Yukhi and Stephen J.
O’Brien, Laboratory of Viral Carcinogenesis, National
Cancer Research Institute, Bldg. 560 Room 21- 105,
Frederick, MD 21702-1201, U.S.A.

Annual budget: III
Time frame: 5 years
Contact: A.J.T. Johnsingh, Wildlife Institute of India, P.O.
Box 18, Chandrabani, Dehra Dun 248 001, Uttar Pradesh,
India.

70. Conservationof the Iriomote cat*Objective: To ensure the conservation of viable nu.mbersof Iriomote cats in the wild.

Jungle cat (Fe/is chaus): Category 4
No projects suggested. Some information on distribution
will be collected during surveys for other sympatricspecies in certain parts of the jungle cat’s range (Projects
59,63, 65).

Description: Conservation of the Iriomote cat is politicallydifficult because most cats occur on the coastal lowlands,
rather than the mountainous center of the island where theprotected area is, and suffer a high degree of humanrelated mortality, including road kills. Study of the ecology and population dynamics of the Iriomote cat,sponsored mainly by the Japanese government and continuing now for more than a decade, will shift to the so-far
unstudied western portion of the island, where prospects
for lowland reserve creation are best due to its relatively
low level of development.

Leopard cat (Prionailurus
bengalensis):
Category 5
See Project 13 under General Topics, Trade. Distribution
information will be collected by Projects 59, 62, 63, and65.

lriomote cat (P.6. iriomotensis):but high priority

Plan

Annual budget: V

Not ranked,

Timeframe: 5 years
69. Evaluation of the taxonomic
status of the Iriomote cat*Contact: Masako Izawa, Dept. Biology, University of the
Ryukus, Nishihara, Okinawa 903-O 1, Japan.

Objective: To determine the taxonomic status of the
Iriomote cat by reconciling the opinions of different
experts in the fields of systematics, anatomy, genetics,
and ethology.

Eurasia
Asia sub-region

Description: The Iriomote cat is found only on the small
Japanese island of Iriomote, the southernmost island in
the Ryuku chain, located about 200 km east of Taiwan. It
has variously been considered a monotypic species(Mayailurus iriomotensis); a species closely related to theleopard cat (Prionailurus iriomotensis); and a subspeciesof the leopard cat (P. bengalensis iriomotensis). The project executant will liase with the various experts on the
Iriomote cat with the aim of producing a consensus decision on how the cat should be scientifically classified. The
executant will also work with experts in the Philippines tocompare the degree of differentiation of these island populations. The classification of the Iriomote cat has important repercussions for conservation action: is it the world’s
most endangered cat, or a distinctive island population of
one of the world’s more common cat species?

Snowleopard(Uncia uncia): Category

1 (A)71. Support for the International
Snow Leopard Trust*Objective: To support the activities of Project Snow
Leopard of the International Snow Leopard Trust inCentral Asian range states.
Description: The snow leopard ranks with the tiger as themost vulnerable of the big cats. The population isextremely fragmented, and snow leopards are threatenedby persecution for livestock predation and by commercial
poaching. The International Snow Leopard Trust hasdeveloped a substantial body of expertise on the species
since its establishment in 1981. It is developing a coordinated information program to monitor the status of snow
leopard populations, and to develop a cooperative frame-

Annual budget: II
Time frame: 1-2 years

295Part III: Cat ActionPlanprotected areas, such as the Changtang and Qomolangma
Nature Reserves, and forest department staff provided withcold weather gear and more reliable transport than currently exists. The project plans to incorporate village-level
wildlife stewardship and is being developed in Nepal(Project 74).

work within which range states can work together. The
program, called “Project Snow Leopard: Conservation ofCentral Asian Biodiversity,”
has six components: (1)involvement of local people in all stages of snow leopard
projects; (2) improving management of mountain reservesin snow leopard range states; (3) strengthening local conservation institutions through training and program support; (4) reducing poaching by providing technicalsupport; (5) promoting regional cooperation; and (6) identifying and establishing habitat corridors and trans-boundary reserves.

Annual budget: I-III
Time frame: 5 years
Contact: Rodney Jackson,
Sonoma, CA 95476, U.S.A.

Annual budget: I-IV (examples: I. development of conservation education materials and dissemination to schools
and villages; II. field training workshops for rangers fromsnow leopard reserves; III. identification of “hotspots” forsnow leopard livestock predation, and development of programs to reduce people-wildlife conflicts; IV. identification and protection of habitat corridors.)

18030 Comstock

Ave.,

73. Evaluation of the status of the snow leopard
in Russia and the central Asian republics*
Objectives: To continue assessment of the snow leopard
populations of Russia and the central Asian republics; andto produce regionally-based conservation management
plans.

Time frame: OngoingContact: Helen Freeman, President, International Snow
Leopard Trust, 4649 Sunnyside Ave. No., Seattle, WA
98103, U.S.A.

Description: The northernmost populations of snow leopard may be sufficiently isolated from the southern populations to warrant taxonomic differentiation; moreover, they
are unique in their occupation of coniferous forest habitat.
Mapping of population distribution needs to be completed,
as well as evaluation of migration possibilities and identification of key habitat corridors. The size of each regional
population will be estimated; the possibility of migrationexchange between them investigated; and limiting factors
identified. Animals will be captured and sampled forgenetic analysis. Preliminary field surveys have alreadybeen carried out in the Altay, Sayan, and Tuva mountain
ranges in southern Siberia, and a detailed ecological studyof snow leopards in the western Tian Shan (Kyrgyzstan)
has been completed. These areas will be further studied
in the first phase of the project. The second phase will
move on to the Dzhungarian Alatau in Kazakhstan, and thePamir and Pamir-Altai in Uzbekistan and Tajikistan.

72. Status and management ofsnow leopards in Tibet*
Objectives: To determine the status of snow leopards insoutheastern Tibet and adjoining areas of China; and to
implement measures aimed at reducing illegal trade inbones and furs.
Description: Tibet probably holds the world’s largest population of snow leopards. Recent work by G. Schaller, R.
Jackson and J. Fox, however, has shown that snow leopard
range is considerably more fragmented than previously
supposed, with large areas containing very low numbers,
or none at all. No status surveys have been conducted insoutheastern Tibet, nor have rumors of snow leopard
occurrence (along with blue sheep) in Yunnan province
and the extreme northern tip of Myanmar been investigated. In the last few years, both ungulate and snow leopard populations have come under increasing poachingpressure. New roads provide relatively easy access touninhabited terrain, but the vast tracts of land makepatrolling extremely difficult. There is an urgent need totrain Tibetans and others in survey and habitat assessment
techniques, so that key wildlife areas can be identified andprotected. Check-posts need to be established in and near

Annual budget: II
Time .frame: 3 years
Contact: Eugene Koshkarev, Tchaikovsky
Street 140,
Apartment 6, Irkutsk, Russia. U.S. address: c/o International Snow Leopard Trust, 4649 Sunnyside Ave. No.,
Seattle, WA 98103.

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Part 111: Cat Action

74. Reducing livestock depredation in the
Annapurna ConservMion Area of Nepal:
implementation of a snow leopard
management plan*

Description: Pakistan holds a small, threatened populationof snow leopards, with numbers estimated at 100-200.
WWF-Pakistan has initiated some small-scale projects insnow leopard areas which have proved successful. Future
activities will include the following: (1) presence/absence
surveys to map out snow leopard population distribution;(2) working with grazier communities to improve antipredator measures; (3) small-scale development programs
which enhance crop and livestock productivity to help offset losses to predators; (4) conservation education programs about the snow leopard for both local graziers andthe general public; (5) training seminars for local wildlife
officials to improve snow leopard protection and enforcement of the trade ban.

Objectives: To investigate factors contributing to livestock
depredation by snow leopards and leopards in three
regions of Nepal’s Annapuma Conservation Area (PhaseI), and to develop and implement measures to reduce lossof livestock and alleviate related people-wildlife conflict(Phase II).
Description:
The recently established
Annapurna
Conservation Area holds what is probably the country’s
largest population of snow leopards. Predation by snow
leopards on domestic livestock owned by people living in
the reserve has been investigated by this project (Phase I:
1991-1993), funded in part by the U.S. Agency forInternational Development.
A number of factors were
found to be statistically associated with livestock predation, including inadequate or lack of daytime guarding;close proximity to a cliff; and grazing alongside a welldefined habitat edge, such as pasture adjoining brushy
areas, broken terrain, or a valley “bowl.” A number of recommendations were generated for reducing livestock
losses by the project, and Phase II will focus upon educating herders on improving their livestock herding andguarding practices, and upon establishing the importanceof snow leopards as a tourist attraction-a
valuable asset
rather than a burden to the local people. The project is partof the Annapurna Conservation Area Project run by theKing Mahendra Trust for Nature Conservation.
Thislarger project seeks to empower local people in management and conservation of natural resources and wildlife,
primarily through eco-tourism.

Annual budget: III
Time frame: 5 years
Contact: Ashiq Ahmad, Conservation Director, WWFPakistan,
Departmentof Environmental
Studies,
University of Peshawar, Pakistan.

Chinesemountaincat (Fe/b

Met!): Category1

76. Natural history, distribution, andstatus of the Chinese mountain cat
Objectives: To gather the first information on the behavior and ecology of the Chinese mountain cat through radiotelemetry; and to conduct presence/absence surveys togain better understanding of its distribution and status.
Description:
A study site in the vicinity of Xining(Qinghai province), where Chinese mountain cats havebeen collected in the past for the Xining Zoo, will beselected. Animals will be captured, radio-collared, andsampled for genetic analysis. The study will examine thenatural history of the species, with emphasis on determining habitat suitability to predict presence/absence and help
with surveys of the cat’s distribution. Such surveys should
be conducted in montane regions along the northeasternedge of the Tibetan Plateau. The Xinjiang cat survey(Project SO) will help determine whether the Chinese
mountain cat occurs west of its very small known range(see Species Account).

Annual budget: IV
Timeframe:Plan

2 years

Contact: Chandra Gurung, Member Secretary, King
Mahendra Trust for Nature Conservation, P.O. Box 37 12.
Kathmandu, Nepal. Rodney Jackson, 18030 Comstock
Ave., Sonoma, CA 95476, U.S.A. Gary Ahlborn, BioSystems Analysis, 3 152 Paradise Dr. Bldg. 39, Tiburon,
CA 94920, U.S.A.

Annual budget; I-II

75. Conservation of snow leopards andtheir habitat in northern Pakistan*Time frame: 2-3 years
Objectives: To evolve and implement a management strategy for the protection of snow leopard from persecution bygrazier communities, and to stop illegal trade of its pelts.

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Plan

Manul (Otocolobus

manul): . Category

surveys will be conducted in selected areas of the Caspian
region, and the results will form the basis of a detailed
population distribution map.

2

77. Natural history of the manul

Annual budget: I-II

Objectives: To gather the first information on the behavior and ecology of the manul through radio-telemetry.

Time frame: 2-3 years
Description: The natural history of the manul has neverbeen studied using radio-telemetry;it appears to bestrongly associated with cold steppe grassland habitats.
A study site will be selected, and animals captured, radiocollared, and sampled for genetic analysis.

Asiatic wildcat (F. silvestris, ornata group):
Category 2
No specific projects suggested. Some data on population
status and distribution will be collected by Projects 42,
76, and 78-80. See also Projects 10 and 15.

Annual budget: I-II
Time frame: 2-3 years

Eurasian

lynx (Lynxlynx): Category3

78. Status of the cheetah, sand cat, andmanul in Baluchistan, Pakistan

80. Status survey for the lynx and othersmall cats in Xinjiang province, China*

Objectives: To conduct surveys to determine the status of
the isolated populations of the manul and sand cat in western Pakistan; and to investigate whether a resident cheetah population is present.

Objectives: To map the distribution and evaluate population status of the cats of Xinjiang province, including theEurasian lynx, manul, Asiatic wildcat, jungle cat, and possibly the Chinese mountain cat.

Description: The montane juniper steppes of Baluchistan
are unusual habitat for the manul, and the small populations found there appear to be isolated from the main population (see Species Account). Presence/absence surveys
will be conducted and site visits made to ascertain population distribution and status. These surveys will also collect data on the sand cat in the nearby Nushki desert region
close to Iran, and potential occurrence of the cheetah.

Description: The arid mountains of central Asia represent
unusual habitat for the Eurasian lynx, and animals here
are traditionally classified as the subspecies din&i. The
status of the lynx and the other cats of Xinjiang is notknown. The project executant will conduct wide-ranging
presence/absence surveys within China’s vast northwestern province, and will focus on investigating the impactof historically high levels of exploitation for furs. Funded
mostly by the Chinese government, this project will generate recommendations for conservation action, including
better control of hunting.

Annual budget: I-II
Time .frame: l-2 years

Annual budget: II
79. Status of the manul in the area of the Caspian Sea

Timeframe: 2 years

Objectives: To conduct presence/absence surveys to determine the status of the manul in Armenia, Azerbaijan, Iran,
Turkmenistan, and Uzbekistan.

Contact: Ablimit Abdukadir,
Xinjiang Institute of
Biology, Pedology, and Desert Research, Urumqi 830 011,
Xinjiang, China.

Description: These countries are home to a reddish form of
the manul which has been termed a separate subspecies,
the Transcaspian manul 0.m. ferrugineus Ognev, 1928.
The status of this taxon has not been examined in detail,but it is described by the regional Red Data Books as“endangered” (Belousova 1993), and has apparently disappeared in recent years from large areas of its historicrange (Bannikov and Sokolov 1984). Presence/absence

Europe sub-region
lberianlynx (Lynx pardinus):

Category

1

81 Implementation of conservation
management measures for the Iberian lynx*

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Parl

Objective: To implement conservation management measures for the largest sub-populations of the Iberian lynx inSpain and Portugal.

III: Cat ActionPlanlinked with the Experimental Center for Captive Breeding;(7) pilot reintroduction program for lynx captured or bornin captivity; (8) monitoring the effects of habitat management and rabbit restocking programees; follow-up of
repopulation of the rabbit and of means of managing habitat; (9) research into alternative means of controlling conflicting predators. This project is affiliated with the LIFE
program of the European Union, and will work closelywith the above Project 81.

Description: The Iberian lynx ranks as the world’s mostvulnerable cat species, reduced to small sub-populations infragmented habitat, with total numbers probably not
greater than 1,200. However, the cat has been well-studied, the threats facing it are fairly well-known, and guidelines for conservation action (ICONA 1992) must now beapplied and tested in the field. These include (1) completion of detailed surveys of the conditions faced by eachlynx sub-population (land use, land ownership, habitat
condition, rabbit density); (2) population viability analysis and genetic sampling; (3) banning rabbit trapping; (4)taking active steps to increase rabbit populations (such asbrush clearance); (5) investigating alternative methods ofcontrolling competing predators of rabbits; and (6) reintroduction-oriented research associated with the captive
breeding/rehabilitation program now underway. This project is affiliated with the LIFE program of the European
Union, and will work closely with the next Project 82.

Annual budget: IV-V
Time frame: 4 years
Contact: Miguel Delibes, Estacion Biologica de Dofiana,
Apartado 1056, Avda. Maria Luisa s/n, 41013 Sevilla,
Spain.

83. Increasing public awareness of theendangered status of the Iberian lynx in Spain*
Objective: To promote conservation of the Iberian lynx inSpain (and reduce inadvertent human-caused mortality)
through a public awareness campaign.

Annual budget: IV-V
Time frame: 4 years

Description: Human-caused mortality, particularly traps
and snares set for rabbits, but also including illegal shooting and road kills, is high for the Iberian lynx in Spain(see Species Account), and can have serious adverse
effects on small isolated populations. This project isdeveloping public education campaigns directed at two
audiences -huntersand school children-to
promoteawareness of the endangered status of the Iberian lynx andthe importance of its conservation. Activities include production and distribution of posters, leaflets, pins, and stickers; travelling lectures; and a children’s art competition
with the lynx as the subject.

Contacts: Spain: Miguel Aymerich, Instituto Nationalpara la Conservation de la Naturaleza (ICONA), Gran Viade San Francisco 4, 28071 Madrid, Spain. Portugal:
Direccao de Services de Conservacao da Natureza,
Ministerio do Ambiente e dos Recrusos Naturais, Institutoda Conservacao da Natureza, Rua da Lapa 73, 1200
Lisboa, Portugal.

82. Applied studies on theconservation of the Iberian lynx*
Objectives: To continue the long-term study of the behavior, ecology, and conservation status of the Iberian lynx inSpain.

Annual budget: II

Description:
The Lynx Study Group, based in Coto
Donana National Park, has produced most of the biological data on the Iberian lynx, as well as training a number ofstudents, and clarifying the conservation status and problems of the most vulnerable of the cat species. Future
study topics include the following: ( 1) population statussurveys; (2) genetic and pathological research; (3) use of
habitat in relation to other predators and development of ageneral habitat model for the lynx; (4) plan for habitat corridors to avoid fragmentation of distribution areas; (5)models for analysis of population viability; (6) researchContact: Sociedad Espafiola para la ConservationyEstudio de 10s Mamiferos (SECEM), Depto. Biologia
Animal, Universidad de Malaga, 2907 1 Mlaga, Spain.

Timeframe:

EurasianI yearlynx (L. lynx): Category2

84. Support for the Status and Conservationof the Alpine Lynx Population Project*
Objectives: To re-establish the lynx in the Alps through

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Parl

III: Cat ActionPlanvores, particularly the wolf and the brown bear, which areexpanding their ranges in the Alps where lynx have been
reintroduced. The researchers will emphasize re-establishing historic connectivity in European lynx populations
through Austria, where nine lynx were released in 19771979, but a population was apparently not established.

international cooperation by (1) defining the present distribution and status of the lynx in the Alps; (2) reviewing
the reintroduction of the lynx into the Alps 20 years after
the first releases; (3) agreeing on methods of monitoring
the development of sub-populations; and (4) proposinginternationally coordinated conservation measures where
necessary.

Annual budget: III
Description: Lynx were eradicated in the Alps during the
19th century. The Alps are one of the last near-naturalareas of central and western Europe, stretching from
France through Switzerland and Italy to eastern Austriaand Slovenia over an area of nearly 200,000 km? Thesemountains could sustain a population of more than 1,000
lynx, and could connect and support smaller potential populations in secondary mountain chains. Lynx have been
reintroduced into the Alps several times since the 1970s(Austria, Germany, Italy, Slovenia, Switzerland) with
mixed results. Most reintroduced populations have notbeen monitored. In recent years, heavy casualties due toillegal killing and road accidents have halted the expansionof at least the Swiss population. Inbreeding is also a potential threat. The long-term goal of the reintroductions must
be to connect the sub-populations into a single Alpine
meta-population. This can only be accomplished throughinternational cooperation.
This project is run by theEurasian Lynx Group of the Cat Specialist Group, and isfunded in part by the Swiss Federal Office of theEnvironment and WWF Switzerland.

Time frame: 3 years
Contact: Thomas Huber, Institut fur Wildbiologie und
Jagdwirtschaft, Universitat fur Bodenkunde, Peter-JordanStrasse 76, A-l 190 Wien, Austria.

86. Preliminary assessment of lynx
status in the Caucasus region*
Objective: To conduct a preliminary assessment of lynx
distribution and statu S in the Caucasus mountains ofArmenia, Azerbaijan, Georgia, and Russia.
Description: The Caucasus mountains are believed to bea significant reservoir of intraspecific diversity: theseheavily spotted animals were once believed to be conspecific with the Iberian lynx (Lynxpardinus),
as discussed
in the Species Account. They are also believed to be veryrare and threatened-a recent WWF survey of Georgia’s
Borzhom Nature Reserve, an important regional protectedarea for lynx, found no sign of lynx presence. This project will conduct further surveys in key areas to define current lynx status.

Annual budget: I
Time .frame: OngoingContact: Urs Breitenmoser and Christine BreitenmoserWursten, Swiss Lynx Project, Villettengassli 4, CH-3074
Muri, Switzerland.

Annual budget: I-II
Time frame: 2 years
Contact: Jason Badrize, Institute of Zoology, 31 Chavchavadze Ave., GE 380 030 Tbilisi, Republic of Georgia.

85. Spatial organization, population dynamics,
and feeding ecology of the reintroduced lynx
population in Slovenia*

87. Biology and ecology of the lynx inthe lowland primeval forest of the
Bialowieza Nature Reserve, Poland*

Objective: To study the ecology and population dynamics
of the reintroduced lynx population in Slovenia.
Description: The Slovenian lynx population has undergone
remarkably rapid range expansion since six animals werereintroduced in 1973. The population has not been studied,and this radio-telemetry project will employ the same
methodology used to st udy the reintroduced Swiss lynx
populations. Four lynx have been radio-collared to date.
The ecology of the lynx in Slovenia will provide important data on the lynx’s relationship with other large cami-

Objectives: To document (1) spatial organization, activity
patterns and migration by lynxes through radio-telemetry;(2) the impact of lynx predation on ungulate populations;and to develop (3) a conservation plan for lynx in Poland.
Description: The Bialowieza lynx population is the lastremaining native population of lynxes in lowland primeval

300

Part III: Cat Action

Plan

Description: Wildcats occur on the nearly 100 islands off
the European, African, and west Asian coasts, but theirtaxonomic status is not known. They are believed to be
most closely related to the Zybica group, but it has also
been suggested that some populations have evolved fromferal domestic cats originally introduced to the islands byhumans. This project will address the following problems:(1) present occurrence and distribution; (2) past occurrence
and/or way of colonization; (3) taxonomic and genetic status; (4) natural history; (5) population status; (6) conservation and management problems; and (7) conservationand management strategies. Preliminary work has begun
in Greece, Sardinia, and Corsica. This project will be complementary to Projects 10, 15, and 45.

European forest. Indigenous lynx populations have not yet
been studied through radio-telemetry.
Preliminary studies suggest unusually high levels of predation on red deer.
The ecology of the Bialowieza population will be intensively studied through radio-telemetry, snow tracking,recording of ungulate kills and surveys of ungulate preydensity. At the same time, educational materials will be
produced to help conserve lynx populations in other partsof Poland.
Annual budget: II
Time frame: 3 years
Contact: Henryk Okarma, Mammal Research Institute,
17-230 Bialowieza, Poland.

Annual budget: IV
Time frame: 5 years

88. Ecology and status of lynx
populations in Scandinavia*

Contact: Bernardino Ragni, Instituto de Zoologia, Facolta
di Scienze, Universita degli Studi di Perugia, Via Elce diSotto, Perugia 06 100, Italy.

Objectives: To study the ecology and status of lynx populations in northwestern Europe through radio-telemetry.

90. Status and distribution

Description: The Scandinavian lynx populations are thelargest in western Europe, but their ecology and limiting
factors have not yet been studied with radio-telemetry.
Findings of these projects, which focus on estimating
national populations, mapping their distribution, and identifying limiting factors, will have important implications
for hunting management in the region.

Objectives: To evaluate the status and distribution of wildcat populations in France.
Description: France is home to one of the largest wildcat
populations in Europe. This project, sponsored in part by
the Office National de la Chasse of the Minis&edel’Environnement, will conduct presence/absence surveys
to complete a detailed population distribution map, andevaluate the degree of hybridization with domestic cats.

Annual budget: III
Annual budget: 5 years

Annual budget: I

Contact: Sweden: Tommy Kruger, Dept. of Zoology,
University of Stockholm, Svante Arrhenius vag 14- 16, S106 91 Stockholm, Sweden. Norway: Tvor Kvam, Norsk
Institutt for Naturforskning,
Tungasletta 2, N-7005
Trondheim, Norway.

EuropeanCategory

wildcat3(f. silvestris,

silvestrisof the wildcat in France*

Time frame: 2 years
Contact: Philippe Stahl, Office National de la Chasse,
Grange neuve, 01330 Villars les Dombes, France.

group):The Americas

89. Status survey and taxonomic evaluationof the wildcats of the Mediterranean islands*

Kodkod(Oncifelis

guigna):Category

91. Natural history, distribution,
status of the kodkod*

Objectives: To evaluate the conservation and taxonomic
status of wildcat populations on selected Mediterranean
islands.

1

andObjectives: To conduct the first study of the behavior andecology of the kodkod through radio-telemetry; to collect

301

Part III: Cat ActionPlanrange of the Andean mountain cat. This project will conduct surveys in selected parts of the high Andes to examine: (1) species occurrence; (2) degree of population
isolation; and (3) habitat and prey association variables.
Data will also be gathered on the occurrence of sympatriccat species. If possible, biological samples will be gathered for morphological and genetic analysis. The survey
will permit selection of an appropriate site for the natural
history study component of the project. Baseline behavioral and ecological data will complement and complete
the information gathered by the first component on species
distribution and status.

biological samples for genetic analysis; and to conduct
presence/absence surveys in key parts of its range to map
population distribution and evaluate species status.
Description:
The kodkod has an unusually restricted
range, and is moreover separated into two major population groups: the largest associated with temperate rain forest in Chile and Argentina, and the other with coastal
shrubland in central Chile. The natural history study will
be carried out in the moist temperate mixed forests ofArgentina’s Nahuel Huapi National Park, a habitat typicalof the largest population group, and will greatly advanceunderstanding of the species biology, ecology and status.
Biological samples will be collected for genetic analysis tobetter evaluate taxonomic status, particularly with regard
to Geoffroy’s cat. Finally, the project will contact the
Corporation National Forestal (CONAF) in Chile to conduct presence/absence surveys for the kodkod in bothmajor reserves and in modified habitats, such as farmland
and logged areas, with priority going to central Chile. The
project has support from the Delegation Tecnica Regional
Patagonia of the Administrationde Parques Nacionales
of Argentina (complete radio-telemetry equipment) and ispartially sponsored by the Sociedad Naturalista Andino
Patagonica.

Annual budget: II
Time frame: 3 -4 years

JaguarCategory2(A)Objectives: To examine cattle mortality on ranches in theBrazilian Pantanal, determine what percentage is attributable to large cats, and develop specific management recommendations for big cats in the region.
Description: The seasonally flooded inland Pantanal deltaarea, the largest of its kind in the world, protects an important population of jaguars, as well as pumas. Jaguar distribution has been mapped in the Pantanal (most occur
outside of protected areas: Quigley and Crawshaw 1992),
and the percentage of livestock in jaguar and puma diets
living on one ranch has been quantified (Crawshaw andQuigley 199 1 and in prep.). This study will complement
the others by examining overall cattle mortality throughmonitoring of ranch records; accompanying ranch hands
on patrols; cattle carcass collection; and by quantifying
calf suvival by fitting 50-60 calves with radio collars
equipped with mortality sensors. Jaguars and pumas will
also be radio-collared. Data obtained on movement, activity and predation will be analyzed in relation to cattle management practices to evaluate the impact of these predators
on ranch livestock. Results of the study will be used todevelop a big cat conservation management plan for theregion, suggesting alternative practices which may reduce
economic losses resulting from depredation. The study
area encompasses both a high-density cattle ranch and theadjacent 1,400 km2 Pantanal National Park.

Time .frame: 2-3 years
Contact: Osvaldo Nestor Herrera, c/o Claudio Chehebar,
Delegation Tecnica, Regional Patagonia, Administrationde Parques Nacionales, Intendencia de1 Parque National
Nahuel Huapi, 8400 San Carlos de Bariloche, Rio Negro,
Argentina.

cat (Oreaihwsonca):

93. Jaguar and puma depredation of livestockin the Pantanal of Mato Grosso, Brazil*

Annual budget: II

Andean mountain
Category 1(Panther-ajacobitus):

92. Natural history, distribution, andstatus of the Andean mountain cat
Objectives: To gather the first information on the behavior and ecology of the Andean mountain cat through radiotelemetry; to collect biological samples for genetic
analysis; and to conduct presence/absence surveys in thehigh Andes to map population distribution and evaluate
species status.
Description: The Andean mountain cat, with a restricted
range in the high alpine zones of the Andes mountains ofSouth America, is one of the least known cats in the world.
There are few existing records by which to document theAnnual budget: IV
Time frame: 3 years

302

Part III: Cat ActionPlanof radio-telemetry; jaguar and prey densities estimated in
the reserve; and a conservation management plan will be
produced.

Contact: Peter Crawshaw, IBAMA, Projeto Camivoros,
Parque National do Iguacu, C.P. 750, Foz do Iguacu,
Parana 85 85 l-970, Brazil.

Annual budget: V
94. Ecology of jaguars and othercarnivores in the Brazilian cerrado*

Time frame: 3 years
Contact: John W. Laundre, Department of Biological
Sciences, Idaho State University, Pocatello, ID 83209,
U.S.A.

Objective: To study, through radio-telemetry, the ecology
of a carnivore community in the Brazilian cerrado.
Description: The cerrado (Savannah woodland) is the second largest habitat type in Brazil (after tropical rain forest),
but the ecology of the cats and other carnivores living there
has never been studied. Emas National Park is the largest
protected area (1,3 10 km*) in the cerrado, and is potentially home to six felid species: jaguar, puma, margay,
ocelot, pampas cat, and jaguarundi. This project willattempt to capture and radio-collar a representative sampleof each species, along with other carnivores, and will compile the first scientific overview of the ecology of a cerradocarnivore community. Its results will aid assessment ofspecies status over a large area of central Brazil, particularly significant for the jaguar.

See also Project 6.

OncillaHgrinus):Category2

96. Natural history of the oncilla,
margay, and ocelot
Objectives: To conduct a comparative study of the behavior and ecology of the Leopardus species in an area where
they occur sympatrically.
Description: This project will be the first to study the natural history of the oncilla in any detail, and will also examine ecological niche partitioning with its close sympatricrelatives, the margay and the ocelot. It should yield datathat will help further understanding of the oncilla’s distribution, especially association with lowland tropical rain
forest in the Amazon basin (see Species Account andProject 97).

Annual budget: III
Timeframe:(Leopardus

5 years

Contact: Peter Crawshaw, IBAMA, Parque National do
Iguacu, C.P. 750, Foz do Iguacu, Parana 85851-970,
Brazil. Leandro Silveira, Depto. de Ciencias Biologicas e
Biomedicas, Universidade Catolica de Goias, Avenida
Universitaria 1440, Setor Universitario, Goiania, Goias
742 10, Brazil.

Annual budget: IV
Time frame: 4-5 years.

95. Conservation of the jaguar in
the tropical dry forests of Mexico*

97. Distributionof the oncilla

Objectives: To map the distribution
selected parts of its range.

Objectives: To study the natural history of a northern population of jaguars in tropical dry forest habitat; and to evaluate the potential of the proposed Chamela-Cuixmala
Biosphere Reserve to support a viable jaguar population.

of the oncilla in

Description: The oncilla’s distribution is little known. It
appears to be strongly associated with montane cloud forest, but has been recorded from other habitat types, including tropical dry forest, subtropical forest, and eucalyptus
plantations. It appears to be absent from much of the lowland tropical rain forest of the Amazon basin, a strongholdfor its close sympatric relatives, the ocelot and the margay.
A Cat Specialist Group regional coordinator will oversee
presence/absence surveys for the oncilla in selected parts of
its range to clarify species distribution. Data on sympatricspecies occurrence will also be collected, and survey mate-

Description: Biosphere reserves are designed to preserveadequate areas to maintain viable poulations of plants andanimals. It is often unknown if a biosphere reserve. suchas the proposed Chamela-Cuixmala
reserve in central
coastal Mexico, is adequate for large carnivores. This area
represents one of the northernmost jaguar populations, andthis study will gather data and make recommendations
necessary to ensure its viability. Jaguar ecology, movements, and spatial organization will be studied with the aid

303

Part III: Cat Action

Plan

Objectives: To gather the first information on the behavior and ecology of these two sympatric cat species in the
habitat type that makes up the majority of their ranges; toevaluate ecological niche partitioning between the two
species; and to map population distribution.

rials and techniques will be carefully designed to avoid confusion between the oncilla and the other small spotted cats,on which data will also be collected.
Annual budget: I-II

Description: Pampas grasslands make up a large part of
the ranges of both the pampas cat and Geoffroy’s cat, but
the natural history of these species has not been studied.
For Geoffroy’s cat, there has been one natural history
study in the southernmost part of its range; for the pampas cat, none. This study will evaluate, through a radiotelemetry study and wider-ranging
presence/absence
surveys, ecological separation between the two species,
and their response to cattle ranching and other forms ofhabitat modification. Recommendations for conservationof the small cats of the pampas will be made.

Timeframe: 3 years

Margay (Leopardus
uviedi): Category 3
See Projects 96 and 97 above for the oncilla.

Canadalynx (Lynxcanadensis):

Category

3

98. Long-term studies of the effects ofharvest on northern Canada lynx populations*

Annual budget: III

Objective: To continue two long-term studies of the effects
of fur harv ,esting on northern populations of Canada lynx.

Time frame: 3-4 years
Description: Two provincial government-sponsored projects are underway in northern Canada, the stronghold of
the Canada lynx, to evaluate the effects of trapping on population dynamics. The Yukon study compares a protected
population to a trapped population, with focus on the role
played by natural refugia in population recovery. The
study in the Northwest Territories monitors hare and lynx
densities in an untrapped area. It has been pointed out that
there have been no studies of the dynamics of a Canada
lynx population throughout an entire IO-year hare-lynx
cycle; both of these studies, underway for several years
now, will fill this gap and greatly advance understandingof the unusual ecology of the Canada lynx. In addition,
they will help to shape regulation of trapping in an area
which constitutes the stronghold of Canada lynx range.

Puma (Puma concoloo:

Category4(A)100. Support for investigations into thepresence of pumas in eastern North America*
Objective: To continue investigation into reports of puma
occurrence in eastern North America
Description: Pumas have been considered extinct in eastern North America for close to a century. However, asteady stream of observations points to either (1) remnant
populations; (2) recent recolonizations; or (3) escapes from
captivity. This project, in association with the Friends ofthe Eastern Panther and other local puma conservationgroups, will undertake the following activities: (1) publication of a field guide to pumas and their sign; (2) publication of a bibliography
on eastern pumas; (3)communication and investigation of future puma sightings.
A conference on eastern pumas was held in 1994, and a
newsletter is produced bi-annually.

Annual budget: ITT
Timeframe: 2-5 years
Contact: Brian Slough, Fish and Wildlife Branch, Yukon
Dept. of Renewable Resources, Box 2703, Whitehorse,
YT Y IA 2C6, Canada; Kim G. Poole. Wildlife
Management Division, NWT Renewable Resources,
Yellowknife, NWT Xl A 2L9, Canada.

Annual budget: II
Timeframe: 2 years

Geoff roy’scat (Oncifelis

geoffroy):

Category

Contact: Jay Tischendorf, American Ecological ResearchInstitute, P.O. Box 380, Fort Collins, CO 80522, U.S.A.

3

99. Natural history and distribution of
Geoffroy’s cat and the pampas cat in theArgentinian pampas grasslands

304

Part III: Cat Action

101. Support for Proyecto Puma in Patagonia*

Annual budget: II-III

Objective: To continue the long-term study of the ecology of the puma in Chilean Patagonia.

Time frame: 6 years

Plan

Contact: Marcello Mazzoli, Projeto Puma, Universidade
Federal de Santa Catarina-UFSC,
Campus Universitario,
Laboratorio de Mamiferos Aquaticos, Florianopolis88015600, Santa Catarina, Brazil.

Description: In cooperation with the Chilean Corporation
National Forestal (CONAF), Proyecto Puma has been running in Chile’s Torres de1 Paine National Park for nearlya decade, and has gathered important data on the behavior
and ecology of pumas in the southernmost part of theirrange. The pumas are well-habituated to humans, and park
authorities are increasingly worried about potential harm
to tourists. This will be one aspect of the future activitiesof the project. The other will be to quantify the impact of
puma predation on guanacos, the major ungulate preyspecies of puma in the southern Andes.

103. Long-term study of puma ecology
in southwestern Alberta, Canada*
Objective: To continue the long-term study of pumas in a
major wilderness area of southwestern Canada, withemphasis on predator-prey dynamics, and quantifying thepuma’s prey requirements.

Annual budget: III
Description: Pumas have been studied in the Sheep River
area of southwestern Alberta since 198 1. Major preyspecies are annually censused by the Alberta Fish andWildlife Division, and puma numbers and predation rates
are estimated through daily monitoring of radio-collaredindividuals.

Time frame: 3-5 years
Contact: William L. Franklin, Dept of Animal Ecology,
124 Science II, Iowa State University, Ames, IA 500 ll322 1, U.S.A.

Annual budget: III
102. Projeto Puma: conservation of the pumain a densely settled region of southern Brazil*Time frame: OngoingContact: Martin Jalkotzy and Ian Ross, AssociatedResource Consultants (ARC), 2201 34th Street SW,
Calgary T3E 2W2, Alberta, Canada.

Objectives: To map the distribution of the puma in thesouthern Brazilian states of Rio Grande do Sul and Santa
Catarina, and to work with farmers to develop techniques
to limit livestock losses to pumas.

See also Project 6.
Description: Southern Brazil is relatively densely settled,
and the jaguar has almost disappeared from this region.
Pumas still exist in fragmented montane habitat, but persecution in response to livestock depredation could lead totheir extirpation. The NGO, Projeto Puma, which is sponsored in part by the Brazilian government, Instituto
Brasileiro de Meio Ambiente e dos Recursos Naturais
Renovveis (IBAMA), is studying the behavior of pumasin this area, their response to habitat encroachment andhuman interference, and the extent and nature of livestock
depredation incidents. The project is relevant to big cat
conservation in settled areas. The first component, partly
completed, is a short-term comparative study of predation
rates between farms with different management techniques, which aims to identify the most important factors
affecting livestock loss. The second component will be
testing the effectiveness of various measures to reduce
depredation, including wild prey population supplementation, different forms of herd management, and the use ofnauseating substances on puma-killed livestock carcasses.

Ocelot (Leopardus
pardalis):
Category 4
See Projects 96 and 97. No other specific projects suggested.

Bobcat(Lynx

rufus):

Category

4104. Distribution and status of thebobcat and puma in the Mexican Sierras
Objectives: To map the distribution of bobcat and puma
populations in selected areas of Mexico and assess species
status.
Description: The dry scrub, oak, and pine forest habitatsof the Mexican Sierras have suffered the highest degreeof transformation and degradation relative to other habitat types, and dryland scrub habitat is under-represented in

305

Part III: Cat ActionPlanimens from museums around the world, suggests that the
pampas cat may be actually composed of three species(see Species Account). Genetic analysis is needed to test
these results, which would obviously have important
implications for species status and prioritization of conservation action.

the system of protected areas (Flores-Villela
andFernandez 1989). Surveys will be carried out in selectedareas to map the distribution of the bobcat and the puma inthese habitat types, and assess the status of key populations. This area represents the southernmost part of bobcat range and, for pumas, links South and North American
populations.

Annual budget: III
Annual budget: I-II
Time frame: 2 years
Time frame: 2 years

Pampascat (Oncifelis

colocolo):Category

Contact.- Dr. Rosa Garcia-Perea, Museo National deCiencias Naturales, Jose Gutierrez Abascal 2, 28006
Madrid, Spain.

4105. Evaluation of the taxonomic
status of the pampas cat*

See also Projects 94 and 99 above. Additional data on natural history and distribution in the high Andes will be gathered by Project 92.

Objectives: To determine, through morphological andgenetic analysis, whether the “pampas cat” is actuallymore than one species.

Jaguarundi(Herpailurusyaguarondi):
Category 5
No specific projects suggested. Some data on natural history and distribution will be gathered in the course of surveys for sympatric cat species (Projects 94 and 97).

Description: Results from the first component of this project, morphological analysis of nearly 80 pampas cat spec-

306

Appendix

Appendix

1. Cat Subspecies

1

Classically Described Cat Subspecies
Derived from Compilations

Family Felidae,
Subfamily

Acinonyx

G. Fischer, 1817

Acinonychinae

Brookes,by C. Groves

Pocock,Catopuma

1917

Caracal1858.

Catopuma temmincki (Vigors and Horsfield, 1827).
Asiatic golden cat.
dominicanorum Sclater, 1898. Southern China.
temmincki Vigors and Horsfield, 1827. Sumatra north tosub-Himalayan region. Probably there is more than
one recognizable subspecies in this region.
tristis Milne-Edwards, 1872. Highlands of southwestern
China, where the spotted form of the golden cat ismost prominent.

1828.

Felinae Fischer,

Severtzov,

Catopuma badiu (Gray, 1874). Bornean bay cat.
No subspecies described.

Acinonyx jubatus (Schreber, 1775). Cheetah.
hecki Hilzheimer, 19 13. Northwest Africa.
jearsoni Smith, 1834. East Africa. Probably includes
ngorongorensis and obergi Hilzheimer, 19 13; andraineyi and velox Heller, 19 13.
jubatus Schreber, 1775. Southern Africa.
soemmerringi Fitzinger, 1855. Somalia and Eritrea toLake Chad.
venaticus Griffith, 182 1. Arabia to central India, now
found only in Iran, includes raddei Hilzheimer, 19 13.

Subfamily

and A. Shoemaker

Fe/is Linnaeus,

1758.

Felis bieti Milne-Edwards,
1892.
Chinese mountain cat.
The forms chutuchta Birula, 1917 and veZZerosa Pocock,1943, described from the Gobi Desert region, probablyrefer respectively to an Asian desert wildcat and adomestic cat (Groves 1980). No subspecies recognized.

1817

Felis chaus Schreber, 1777. Jungle cat.
afinis Gray, 1830. Sub-Himalayan region.
chaus Schreber, 1777. Jordan Valley and Mesopotamia
north to the Caucasus Mountains region and east
through the deserts of the Caspian and Aral seas.
Includes furax de Winton, 1898 and oxiana Heptner,
1969.
@vidina Thomas, 1928. Vietnam. Possibly includes
Burma and Thailand, or these may represent an
undescribed race.
kelaarti Pocock, 1939. Southern India and Sri Lanka.
kutas Pearson, 1832. Northern India and Pakistan,
includes prateri Pocock, 1939.
nilotica de Winton, 1898. Lower Nile River valley,
EgYPtaGray, 1843.

Caracal caracal (Schreber, 1776). Caracal.
algirus Wagner, 1841. Maghreb region, Morocco
to Algeria.
caracal Schreber, 1776. South Africa.
damarensis Roberts, 1926. Namibia.
limpopoensis Roberts, 1926. Northern Transvaal.
Zucani Rochebrune, 1885. Grasslands of southeastern
Gabon.
michaelis Heptner, 1945. Deserts of the Caspian Sea
region, east to the Amur Darya River.
nubicus Fischer, 1829. East Africa, north to the NubianDesert and west to Cameroon.
poecilotis Thomas and Hinton, 1921. Nigeria.
schmitzi Matschie, 19 12. Palestine east to India.

FeZis margarita Loche, 1858. Sand cat.
harrisoni Hemmer, Grubb and Groves, 1976. Arabia.

307

Appendix

1. Cat Subspecies

sizvestris Schreber, 1775. Europe, east to the Carpathian
mountains and the river Dnieper north of the BlackSea.

margarita Loche, 1858. Sahara, includes meinertzhageni
and airensis Pocock 1938.’
schefili Hemmer, 1974. Pakistan, perhaps west to Iran.
thinobia Ognev, 1927. Deserts east of the Caspian sea.

Named island subspecies include: cretensis Haltenorth,1953 (Crete); jordansi Schwarz, 1930 (Balearic islands);
and reyi Lavauden, 1929 (Corsica).

Felis nigripes Burchell, 1824. Black-footed cat.
nigripes Burchell, 1824. Namibia through the Kalahari
to the northern Transvaal.
thomasi Shortridge, 193 1. Eastern Cape Province,
South Africa.
Felis silvestris Schreber,

Herpailurus

Severtzov,

1858.

Herpailurus yaguarondi Lacepltde, 1809. Jaguarundi.
ameghinoi Holmberg, 1898. Western Argentina, southeast to Rio Negro province.
cacomitli Berlandier, 1859. South Texas and neighboring regions of northeastern Mexico.
eyra Fischer, 1814. Southern Brazil through Paraguay tonorthern Argentina.
fossata Mearns, 1901. Honduras to southern Mexico.
melantho Thomas, 1914. Andean valleys of Peru andupper Amazonia, Brazil.
panamensis Allen, 1904. Ecuador through western
Colombia to Costa Rica.
tolteca Thomas, 1898. Sinaloa, Mexico to Arizona.
yagouaroundi Geoffroy, 1803. Amazon basin of Brazil
north to the Guiana highlands.

1775. Wildcat.

Zybica groupbrockmani Pocock, 1944. Somalia.
cafra Desmarest, 1822. South Africa, Zimbabwe,southern Mozambique.
foxi Pocock, 1944. Bush country from Senegal toLake Chad.
griseEda Thomas, 1926. Kalahari region to southern
Angola.
iraki Cheesman, 1920. Arabian desert regions.
gordoni Harrison, 1968. Batinah coast of Oman.
ZybicaForster, 1780. Desert regions of North Africa to
Sudan and northern Niger; probably includes haussa
Thomas and Hinton 1920, Zowei and Zvnesi
Pocock,d
1944.
mellandi Schwann, 1904. South-Central Africa (southern
Zaire, Zambia, Malawi, northern Mozambique).
nesterovi Birula, 19 16. Mesopotamian region to southwestern Iran.
ocreata Gmelin, 179 1. Ethiopian highlands.
pyrrhus
Pocock, 1944. Northern Angola and south*
western Zaire.
sarda Lataste, 1885. Coastal Maghreb region (Morocco
and Algeria) and Sardinia (probably introduced).
tristrami Pocock, 1944. Palestine and Red Sea coastof Arabia.
ugandae Schwann, 1904. East Africa, includes nandae
and taitae Heller, 19 13.

Leopardus

Gray, 1842.

ornata groupcaudata Gray, 1874. Deserts east of the Caspian Sea
to the Tian Shan Mountains, includes matschiei andmurgabensis Zukowsky,
19 14.
ornata Gray, 1830. India. Probably east through Iran.
shawiana Blanford, 1876. Xinjiang and Mongolia,
includes chutuchta Birula, 19 17 and kozlovi Satunin,
1905.

Leopardus pardalis (Linnaeus, 1758). Ocelot.
aequatorialis Mearns, 1902. Northern Andes.
albescens Pucheran, 1855. Northeastern Mexico to
Texas and, historically, Lousiana.
maripensis J.A. Allen, 1904. Venezuela east to Guiana
highlands.
mitis Cuvier, 1820. Southern Brazil through Paraguay
to northern Argentina.
nelsoni Goldman, 1925. Western Mexico from Oaxaca
to Sinaloa.
pardaZis Linnaeus, 1758. Southern Mexico throughCentral America, includes mearnsi J.A. Allen, 1904.
pseudopardalis Boitard, 1842. Northern Colombia andwestern Venzuela.
pusaeus Thomas, 19 14. Coastal Ecuador and perhaps
Yeru.
sonoriensis Goldman, 1925. Northwestern Mexico
to Arizona.
steinbachi Pocock, 194 1. Bolivian highlands.

sihestris group
caucasica Satun in, 1905. Caucasus mountains andTurkey.
grampia Miller, 1907. Britain, now restricted to
Scotland.

Leopardus tigrinus (Schreber, 1775). Oncilla.
oncilla Thomas, 1903. Costa Rica.
pardinoides Gray, 1867. Northern Andes.
tigrinus Schreber, 1775. Northeastern Brazil and
Guyanas.
308

Appendix

Lynx Kerr, 1792.

Leopardus wiedi (Schinz, 1821). Margay.
amazonicus Cabrera, 19 17. Upper Amazon.
bohviae Pocock, 194 1. Andean slopes.
cooperi Goldman, 1943. Southeastern Texas andadjoining regions of Mexico.
glauculus Thomas, 1903. Dry country of Mexico; probably includes oaxacensis Nelson and Goldman, 193 1.
nicaraguae J.A.Allen, 19 19. Central America; probably
includes salvinius Pocock, 194 1.
vigens Thomas, 1904. Northeastern Brazil to the
Guyanas.
wiedi Schinz, 1821. Southeastern Brazil to northeastern
Argentina.
yucatanicus Nelson and Goldman, 193 1. Rainforest
regions of Mexico.

Leptailurus

Severtzov,

1. Cat Subspecies

Lynx canadensis Kerr, 1792. Canada lynx.
canadensis Kerr, 1792. Mainland northern U.S. andCanada.
subsolanus Bangs, 1897. Newfoundland island.
Lynx lynx (Linnaeus, 1758). Eurasian lynx.
carpathicus Kratochvll and Stollmann, 1963. Carpathian
Mountains west to Bulgaria and Greece.
dinniki Satunin, 1915. Caucasus Mountains south toTurkey and northern Iran.
isabelLinus Blyth, 1847. Kashmir and Tibet north to the
Tian Shan and Altai mountain ranges in Xinjiang andMongolia.
kozlovi Fetisov, 1950. Central Siberia, from the Yenisei
River to Lake Baikal.
Lynx Linnaeus, 1758. Northern and western Europe east
to the Yenisei River in Russia.
neglectus Stroganov, 1962. Russian Far East, Korea, andnortheastern China (Manchuria); includes stroganovi
Heptner, 1969.
wrangeli Ognev, 1928. Eastern Siberia, south to the
Stanovoy mountains.

1858.

Leptailurus serval (Schreber, 1776). Serval.
Many, probably too many, subspecies have been
described. Weigel ( 1961) points out that there is a
general division between a dark, fine-spotted type
(brachyurus and lipostictus) and a lighter, large-spotted
type (most of the rest), with beirae, togoensis, andpococki being somewhat intermediate.

Lynx pardinus (Temminck,
No subspecies described.

brachyurus Wagner, 1841. Sierra Leone. May include
pococki Cabrera, 19 10 (Senegal) and togoensis
Matschie, 1893 (Dahomey gap).
constantinus Forster, 1780. Northern Morocco andAlgeria.
hindei Wroughton, 1910. Kenya, east of the Rift Valley.
May include ferrari de Beaux, 1924 (southern
Somalia).
*
Ziptostictus Pocock, 1907. Uganda, Zaire, and northern
Angola moist forest. May include faradjius Allen,
1924 (northeastern Zaire); kivuensis Lonnberg, 19 19(Lake Kivu region) and kempi Wroughton, 19 10(Kenya west of the Rift Valley).
phillipsi Allen, 19 14. Ethiopian highlands west to
Lake Chad.
serval Schreber, 1776. Southern Zaire and Tanzania,
south to Cape Province. Probably includes beirae
Wroughton, 19 10 (Beira district, Mozambique);hamiltoni Roberts, 193 1 (northern Transvaal); ingridi
Lundholm, 1955 (Zimbabwe); Zimpopoensis Roberts,
1926 (Botswana and western Transvaal); lonnbergi
Cabrera, 1910, mababiensis Roberts, 1932 and niger
Lonnberg, 1897 (southern Angola).
tanae Pocock, 1944. Dry zone of Ethiopia, Eritrea andnorthern Somalia.

1827). Iberian lynx.

Lynx rufus (Schreber, 1776). Bobcat.
After Hall (198 l), with comments by Samson (1979).
Read (198 1), based on differences between over 950
bobcat skulls from the southcentral U.S., considers that
too many subspecies were recognized by Hall, pointing
to the lack of geographic barriers between them.
baileyi Merriam, 1890. Southwestern arid zone from
California to western Texas and Utah, and south toDurango, Mexico.
californicus Meams, 1897. Nevada to central andsouthern California.
escuinapae Allen, 1903. Central Mexico. Samson(1979) suggests that the subspecies is invalid, being
very similar to californicus and texensis; however, he
had a small sample size and did find it to differ frombaileyi, the subspecies directly to the north.
fasciatus Rafinesque, 18 17. Coastal forests from southwestern British Columbia to northern California.
floridanus Rafinesque, 1817. Southeastern U.S.
gigas Bangs, 1897. Maine and adjacent southeastern
Canada, including Nova Scotia.
paEZescens Merriam, 1899. Rocky mountains from
British Columbia to New Mexico.
peninsularis Thomas, 1898. Baja California, Mexico.

309

Appendixc

1. Cat Subspecies

Oncifelis geoffroyi (d’orbigny
and Gervais, 1844).
Geoffroy’s cat.
after Ximenez ( 1975)
euxanthus Pocock, 1940. Bolivian highlands.
geofSroyi d’orbigny and Gervais, 1844. Pampas grasslands from Buenos Aires south to Patagonia.
paraguae Pocock, 1940. Southern Paraguay, southemmost Brazil, Uruguay and northern Argentina.
salinarum Thomas, 1903. Chaco region.

rufus Schreber, 1776. Northeast and central U.S. Samson(1979) recommends that this taxa be split into northeastern and central plains subspecies.
superiorensis Peterson and Downing, 1952. Southeastern
Manitoba to Wisconsin and Minnesota.
texensis, Allen, 1895. Western Lousiana through eastern
Texas to northeastern Mexico.

Oncifelis

Severtzov,

1858.
Oncifelis guigna (Molina, 1782). Kodkod.
guigna Molina, 1782. Southern Chile and Argentina.
tigriZEo Schinz, 1844. Central Chile, includes molinae
Osgood, 1943.

Oncifelis colocolo (Molina, 1782). Pampas cat.
braccatus Cope, 1899. Mato Gross0 to northern
Argentina and the southernmost tip of Brazil.
The pampas cat’s range in Brazil extends throughthe center and into the northeastern region, but
specimens have not been analyzed taxonomically(Silveira in press).
budini Pocock, 194 1. Salta highlands in northwestern
Argentina.
colocolo Molina, 1782. Central Chile.
crespoi Cabrera, 1957. Salta lowlands in northwestern
Argentina.
garleppi Matschie, 19 12. Andes in Peru, Ecuador and
Bolivia.
muiioai Ximenez, 196 1. Uruguay.
pajeros Desmarest, 18 16. Pampas grasslands from
Buenos Aires to southernmost Argentina and Chile.
thomasi Lonnberg, 19 13. Ecuador.

Oreailurus

Cabrera,1940.

Oreailurus jacobitus (Cornalia,
Andean mountain cat.
No subspecies described.

Otocolobus

Brandt,1865).

1842.

Otocolobus manul (Pallas, 1776). Manul.
ferrugineus Ognev, 1928. Kazakhstan south to Iran andPakistan, including the lowlands south of the Caucasus
and west of the Caspian Sea.
man& Pallas, 1776. Lake Baikal region south throughMongolia to Gansu province, China.
nigripectus Hodgson, 1842. Kashmir east to Nepal,
through the Tibetan highlands and east to Sichuan.

Garcia-Perea (1994) considers the pampas cat toconsist of three species, further separated into thefollowing subspecies:
Lynchailurus pajeros (Desmarest, 18 16). Distributed
from the eastern slopes of the Andes in Ecuador south
through lowland habitats to Patagonia. Subspecies:
budini, crespoi, crucinaus Thomas, 190 1 (Patagonia),
garleppi, pajeros, steinbachi Pocock 194 1 (Bolivia),
and thomasi.

Prionailurus

Severtzov,

1858.

Prionailurus bengalensis (Kerr, 1792). Leopard cat.
after Yu and Wozencraft (in press)
alleni Sody, 1949. Hainan Island, off the coast of southem China. Possibly not distinct from chinensis.
bengazensis Kerr, 1792. Indian and Indochinese regions(including China’s Yunnan province), and Malay
peninsula.
borneoensis Brongersma, 1935. Borneo. Possibly not
distinct from sumatranus.
chinensis Gray, 1837. China, except for northeast, andTaiwan.
euptilurus Elliot, 187 1. Manchurian region, Korea andRussian Far East (includes Japanese Tsushima islands
off the coast of South Korea). Heptner (197 1) considered this taxa a separate species, but there are no geographic barriers which would serve to isolate it.

Lvnchailurus
braccatus (Cope, 1899). Distributed
4
through humid, warm grass- and shrubland in Brazil,
Paraguay and Ecuador. Subspecies: braccatus andmufioai.
Lynchailuruscolocolo (Molina, 1782). Distributed asI
two separate populations in Chile. Subspecies: colocola (sub-tropical forests at middle elevations in central Chile) and wo@sohni (Garcia-Perea 1994) on thewestern slope of the Andes in northern Chile.

310

Appendix

Puma Jardine,

hors$&Zdi Gray, 1842. Sub-Himalayan
region east of the.
Indus River.
iriomotensis Imaizumi, 1967. Iriomote Island, Japan.
Some authorities recognize the Iriomote cat as a separate species, Mayailurus or Prionailurus iriomotensis.
javanensis Desmarest, 1816. Java and Bali.
sumatranus Horsfield, 1821. Sumatra and Nias islands.
trevelyni Pocock, 1939. Kashmir.

acrocodia Goldman, 1943. Chaco region of Paraguay
and Boliva.
anthonyi Nelson and Goldman, 193 1. Rain forest ofsouthern Venezuela.
borbensis Nelson and Goldman, 1933. Central
Amazonia.
califomicus May, 1896. California (non-desert, except
far north). Probably includes aztecus Merriam, 1901
(New Mexico and Arizona south to Jalisco); browni
Merriam, 1903 (arid southwestern U.S. to Chihuahua);
and improcera Philipps, 1912 (Baja California,
Mexico).
bangsi Merriam, 1901. Northern Andes (Colombia,
Ecuador, and Peru). Probably includes soderstromi
Lonnberg, 19 13 (northern Ecuador and southern
Colombia).
concolor Linnaeus, 177 1. The Guyanas.
coryi Bangs, 1899. Florida and Lousiana, now restricted
to southern Florida.
costaricensis Merriam, 190 1. Nicaragua to Panama.
Possibly includes mayensis Nelson and Goldman,
1929 (Yucatan peninsula of Mexico south to
Honduras).
cougar Kerr, 1792. Northeastern U.S. and eastern
Canada. Extinct, or nearly so.
greeni Nelson and Goldman, 1929. Eastern Brazil.
hippolestes Merriam, 1897. Rocky mountains in
Wyoming to Idaho and New Mexico. Probably
includes missoulensis Goldman, 1943 (North Dakota
and Yellowstone National Park to Cassiar Mountainsof British Columbia and southwestern Saskatchewan).
hudsoni Cabrera, 1957. Pampas grasslands of Argentina.
kaibabensis Nelson and Goldman, 193 1. Kaibab
plateau, Arizona.
oregonensis Rafinesque, 1832. Coastal forests of British
Columbia south to northern California. Probably
includes Olympus Merriam, 1897 (Olympic mountains,
Washington) and vancouverensis Nelson andGoldman, 1932 (Vancouver Island, British Columbia).
osgoodi Nelson and Goldman, 1929. Bolivian Andes.
pearsoni Thomas, 1901. Type locality Santa Cruzprovince, about 70 miles inland off the coast, southern
Argentina. Synonym probably patagonicus Merriam,
190 1. Type locality Lake Pueyrredon, in the Altiplanoon the border with Chile, Santa Cruz province, southem Argentina (Currier 1983).

Prionailurus plizniceps (Vigors and Horsfield, 1827).
Flat-headed cat.
No subspecies described. Specimens from continental
southeast Asia, Sumatra, and Borneo are slightly
different, but the material is limited.
Prionailurus rubiginosus (I. Geoffroy Saint-Hilaire,1831). Rusty-spotted cat.
phillipsi Pocock, 1939. Wet zone of southwestern
Sri Lanka.
koladivinus Deraniyagala, 1956. Dry zone of Sri Lanka.
rubiginosus I. Geoffroy St.-Hilaire, 183 1. Southern
India. Whether specimens from northwestern India
are of this subspecies is unknown, but there is a skin in
the British Museum of Natural History labelled
Pakistan, which is extremely different.
Prionaihrus
viverrinus (Bennett, 1833). Fishing cat.
rizophoreus Sody, 1936. Java.
viverrinus Bennett, 1833. Sri Lanka, India, continental
southeast Asia and Sumatra.

Severtzov,

1834.

Puma concolor (Linnaeus, 1771). Puma.
Puma subspecies are presently being investigated from agenetic perspective at the laboratory of S.J. O’Brien.

Forms from the Philippine islands remain undescribed.
Rabor (1986) has suggested that the leopard cats ofPanay, Negros, and Cebu, which are separated from theSunda Shelf by deep water channels, may be a different
and endemic subspecies of the Philippines in comparison
with the population found on Palawan, which would be
expected to have a closer relationship to Indonesianisland populations (C. Groves, W. Oliver in Zitt. 1993).

Profelis

1. Cat Subspecies

1858.

Profelis aurata (Temminck, 1827). African golden cat.
aurata Temminck, 1827. West Africa.
cezidogaster Temminck, 1827. Central Africa, east into
Kenya.
Van Mensch and Van Bree (1969) classed all populations
between the Cross (Nigeria) and Zaire (Congo) rivers asintermediate between the two subspecies.

311

Appendix

1. Cat Subspecies

O’Brien et al. (1987b) analyzed samples from captive
African lions and from wild lions of Tanzania’s Serengeti
National Park and South Africa’s Kruger National Park,and suggested that they were of sufficient genetic
similarity to warrant subsummation into a single African
race, P. leo lea.

puma Molina, 1782. Chile across the high cordillera towestern Argentina. According to Cabrera (196 1),
probably includes araucanus Osgood, 1943 (central
Chile); cabrerae Pocock, 1940 (west and central
Argentina); incarum Nelson and Goldman, 1929
(southern Peru); and punensis Housse, 1950
(Tarapaca, Chile).
schorgeri Jackson, 1955. Upper Mississippi River area.
stanleyana Goldman, 193 8. Texas and northeastern
Mexico. S. O’Brien (in CBSG 1991) notes that thistaxa is “genetically similar” to coryi.

Subfamily
Neofelis

Pantherinae

Panthera onca (Linnaeus, 1758). Jaguar.
After Pocock (1939b) and Seymour (1987) and in needof revision, as Pocock’s arrangement was based on very
few specimens and weakly marked distinctions.
arizonensis Goldman, 1932. Originally from Sonora,
Mexico to southwestern U.S.; now extinct in the U.S.
centralis Mearns, 1901. Nicaragua to Colombia.
goldmani Meams, 1901. Yucatan peninsula of Mexico
south to Belize and northern Honduras.
hernandesi Gray, 1857. Tehuantepec and Sinaloa,
Mexico northeast to Lousiana (now extinct in
the U.S.).
onca Linnaeus, 1758. Amazon and Orinoco rain forest.
palustris Ameghino, 1888. Southern Brazil south
through Uruguay to the Rio Negro in Argentina; now
restricted to a few locations in southern Brazil.
paraguensis Hollister, 19 14. Paraguay.
peruviana de Blainville, 1843. Coastal regions of Peru
and Ecuador.
veraecrucis Nelson and Goldman, 1933. Tabasco to central Texas.

Pocock, 1917

Gray, 1867.

Neofelis nebulosa (Griffith, 1821). Clouded leopard.
brachyurus Swinhoe, 1862. Taiwan. Described as a subspecies based upon short tail length, but Pocock(1939) found that this measurement is not a consistentcriterion. Possibly extinct.
diardi Cuvier, 1823. Malay peninsula, Sumatra andBorneo.
macrosceloides Hodgson, 1853. Sub-Himalayan zone
from Nepal to Myanmar.
nebuZosa Griffith, 1821. Indochinese region and southem China.

Panthera

Oken,

Panthera pardus (Linnaeus, 1758). Leopard.
adersi Pocock, 1932. Zanzibar (extinct).
adusta Pocock, 1927. Ethiopian highlands.
ciscaucasica Satunin, 1914. Caucasus mountains.
According to Heptner and Sludskii (1972), the range
is through northern Iran to Afghanistan, and saxicoZor
is a synonym.
dathei Zukowsky,
1964. Southern and central Iran, of
dubious validity.
delacouri Pocock, 1930. Southern China to Malay
peninsula; sinensis is a synonym.
fusca Meyer, 1794. Indian sub-continent.
kotiya Deraniyagala, 1956. Sri Lanka.
japonensis Gray, 1862. North-central China; probably
includes bedfordi Pocock, 1930; chinensis Gray, 1867;
andfontanieri A.M. Edwards, 1867.
jarvisi Pocock, 1932. Sinai peninsula.
leopardus Schreber, 1777. Rain forests of west andcentral Africa; probably includes ituriensis J.A. Allen,
1924.
melanotica Gunther, 1775. Southern Africa; puella andshortridgei Pocock, 1932 are probably synonyms.
melas G. Cuvier, 1809. Java.

1816.

Panthera Zeo (Linnaeus, 1758). Lion.
after Hemmer (1974b).
azandica Allen, 1924. Northeastern Zaire.
bleyenberghi Lonnberg, 19 14. Shaba and Kasai woodland Savannah regions of southern Zaire, and presumably also neighboring parts of Zambia and Angola.
krugeri Roberts, 1929. Kalahari region east to theTransvaal and Natal regions of South Africa.
Zeo Linnaeus, 1758. Coastal woodlands of Morocco
through Tunisia. Extinct.
melanochaita H. Smith, 1842. Cape region, South
Africa. Extinct.
nubica de Blainville, 1843. Northeast and east Africa.
Includes massaica Neumann, 1900; somaliensis
Noack, 189 1; and roosevelti Heller, 19 13.
persica Meyer, 1826. From Iraq to central India in the
19th century; now restricted to Gir Forest, India.
senegalensis Meyer, 1826. West Africa, east to theCentral African Republic.

312

Appendix

I. Cat Subspecies

latter two countries); formerly occasionally ranged
west to Mongolia and Lake Baikal.
amoyensis Hilzheimer, 1905. South-central China, now
restricted to a few locations in southern China.
balica Schwarz, 1912. Bali (extinct).
corbetti Mazak, 1968. Indochinese region north to
Yunnan province, China, and south to the Malay
peninsula.
sondaica Temminck, 1844. Java (extinct).
sumatrae Pocock, 1929. Sumatra.
tigris Linnaeus, 1758. Indian sub-continent.
virgata Illiger, 18 15. Dry river valleys of the Takla
Makan, western slopes of the Tianshan mountains,
Amudarya and Syrdarya river valleys, shores of the
Caspian sea, Elburz mountains, eastern Turkey, Tigris
and Euphrates river valleys. Extinct.

nanopardus Thomas, 1904. Somali arid zone; brockmani
Pocock, 1932 is a synonym, and the taxa probably
includes antinori de Beauz, 1923 (Eritrea).
nimr Hemprich and Ehrenberg, 1833. Arabian peninsula
to southern Israel.
orientalis Schlegel, 1857. Russian Far East, Korea, andnortheastern China; includes villosa Bonhote, 1903.
panthera Schreber, 1777. Coastal woodlands ofMorocco through Tunisia.
pardus Linnaeus, 1758. Sudan and northeastern Zaire;
chui Heller, 19 13 (southern Sudan and Uganda) is
probably a synonym.
pernigra Hodgson, 1863. Kashmir through Nepal tosouthwestern Tibet and Sichuan; synonyms include
millardi Pocock, 1930.
reichenowi Cabrera, 1918. Savannas of Cameroon.
ruwenzorii Camerano, 1906. Ruwenzori and Virunga
mountains of Zaire, Rwanda, and Burundi.
saxicolor Pocock, 1927. Northern Iran and southern
Turkmenia east to Afghanistan.
sindica Pocock, 1930. Southeastern Afghanistan throughwestern and southern Pakistan.
suahelica Neumann, 1900. East Africa, from
Mozambique north to Kenya.
tulliana Valenciennes, 1856. Turkey.

Pardofelis

Severtzov,

1858.

Pardofelis marmorata (Martin, 1837). Marbled cat.
charltoni Gray, 1846. Sub-Himalayan region, from
Nepal to Myanmar.
marmorata Martin, 1837. Continental southeast Asia,
Sumatra and Borneo. Insular and Malay specimens
are, as Pocock pointed out, grey-brown rather thanochraceous like those from the Indochinese region, andsubspecific differentiation is probably warranted.
Moreover, differences between the populations of
Peninsular Malaysia, Sumatra, and Borneo may also be
significant enough to warrant subspecific separation.

Miththapala ( 1992) analyzed subspeciation in the leopard
in terms of genetics. She suggests subsuming all African
races into pardus; all populations on the Indian subcontinent intofusca (i.e., including pernigra); and all
central Asian races into saxicolor. However, only theAfrican samples were considered to be sufficiently
representative. She found the Sri Lankan leopard kotiya
to be well differentiated. Sample sizes were either too
small or inappropriate (coming from captive animals
which were in some cases deliberately inbred or of
uncertain origin) to produce conclusive results fordelacouri, japonensis, melas, and orientalis. She
recommends that her findings be further evaluated byanalysis of pelage.

Uncia Gray, 1854.
Unciu unciu (Schreber, 1775). Snow leopard.
after Stroganov ( 1962)
uncia Schreber, 1775. Northern populations: Central
Asia northeast to Mongolia and Russia.
uncioides Horsfield, 1855. Southern populations: Tibet,
western China, and the Himalayas.

Panthera tigris (Linnaeus, 1758). Tiger.
after Mazak (1979). Genetics are being re-analyzed at
the laboratory of S. O’Brien.
altaica Temminck, 1844. Russian Far East, North Korea,
and northeastern China (possibly extirpated from theAnalysis of subspeciation in the snow leopard is long
overdue; many authorities have suggested that thenaturally fragmented nature of its habitat may have led
to significant differences between populations.

313

Appendix

2. Scientific

Postmortem.

Appendix

Scientific Postmortem:by Andrew2

A Protocol for Collection of Data and Specimens

C. Kitchener,

Steven McOrist, and Robert K. Wayne

iii. Hindfoot length (mm): From ankle to tip of toes,
excluding claws and fur. In the U.S., this measurement includes to the tip of the claws, which is
not relevant for most felids. You may wish tomeasure hind foot length by both methods, whereappropriate.

There is a wide range of important biological informationthat can be obtained from a dead or tranquilized cat. This
protocol is exhaustive, but can be adapted to the needs ofany study. Whenever an opportunity presents itself, specimens should be preserved for taxonomic studies. Other
information would add considerably to our knowledge ofthe basic biology of many species.

iv. Ear length (mm): From notch in base of ear to tipof fleshy part of ear flap. Exclude fur or tufts.
1. External

Featuresf. Teeth: Note number and type, and any damage ordecay, etc., especially if it is not possible to save skull(see below), or if examining a tranquilized cat.

Record the following:a. Color of soft parts: iris, nose, pads, insides of ears.

g. External parasites:
70% alcohol.

b. Coloration and markings of skin: take notes, or color
photographs, of dorsal, ventral, and lateral aspects ofbody, and close-up of face against a standard background.

2. Specimen

d. Weight: if possible, weigh the whole animal. N.B.
Subtract weight of stomach contents (see below), if
these are likely to be a significant proportion of total
weight.
take the following

Preparation

Labelling: All specimens should be labelled. We recommend aluminum tags, on which a number can be
scratched, or embossing type (e.g., Dymo). These canboth be wired on to any skins or skeletal material thatare prepared. The numbers can then be cross-referenced with data in field notebooks. It is recommended
that a second copy be made of this data, in case of accidental loss.

c. Mammae: if female, note number and position of
mammae, and whether lactating.

e. Measurements:
Fig. 1):

Examine fur for these and store inmeasurements (seeb. Scavengers: Watch out for scavengers, whether they
be mammalian, avian, or insect, especially when hanging specimens out to dry. Many valuable specimens
have been lost to scavengers.

Total length (mm): Lay the animal on its back toensure the vertebral column is fully straight and thetip of the nose is in a straight line with the backand the tip of the tail. Never include fur at the tip of
the tail.

c. Skins: If you have time, skin out the specimen, working in fine salt to the newly exposed surface, and also
externally around the ears, eyes, and pads. When fully
skinned, clean off excess flesh and fat, and rub in salt.
If possible, let the skin dry in the sun. When dry, roll
it up and store it somewhere safe and dry.

Tail length (mm): From base of tail dorsally tofleshy tip of tail. Never include fur at the tip of
the tail. It helps to move the tail through 90” tothe body, in order to locate the base of the tail.

In humid habitats where drying may not be possible,
put the skin in a formic acid solution (8 cm3 of 90%

314

Appendix

2. Scientific

Postmorlem

the sun if possible. Store in sealed tins. The worst that
can happen is that the skeleton can get a bit smelly.
e. Where to send specimens: Send specimens to local
universities and museums. If possible, make arrangements with institutions for opportunistic acquisition ofmaterial with respect to arranging import/export
licenses, health licenses, etc. The key to the best useof any specimen is planning before any field work is
started.
Obviously, only rarely will there be time to get all theinformation and specimens outlined above. Those
items in bold lettering are key data and samples which
are required as a minimum.

3. Internal

examination

The purpose of a scientific post mortem is to collect information on biological parameters such as diet, reproductive status, etc., as well as to determine causes of morbidity
and mortality by noting pathological features (i.e., thosethat differ from normal).
To this end, all major organ systems (cardiovascular, respiratory, alimentary, urogenital, and hemolymphatic)should be examined thoroughly. Data and samples should
be examined from a wide range of tissues so that post
mortem investigations are not limited to the one or two
obvious features that may first strike the observer.
Consultation with local veterinary laboratory services
would ensure evaluation of most pathological samples.
Thick slices of all major organs and portions of the gastrointestinal tract should be immersed in 10% buffered formalin. Stomach contents, internal parasites, intestinal
length and reproductive organs should be carefully examined (see below).

Figure 1. Postmortem
measurements.
A = total length in mm; B = tail length in mm; C = hind foot length
in mm; D = ear length in mm.

Where viruses are suspected at a post mortem, samples offresh spleen, liver, and lymph nodes should be collected
and stored chilled at 0” C before being sent to a virus laboratory. Virus transport medium is available at some centers. Blood sampling (see below) of a representative
sample of a population of living cats for virus serology
represents a further effective means of monitoring for thepresence of viruses within a population.

formic acid per liter of water, plus 100 g of salt per literof water). When travelling, the solution can be drained
off and the skins put in plastic bags or dried out before
leaving. Put them in the freezer on arrival.
If desperate, store whole specimen in 70% alcohol oreven methylated spirit. Use syringe to inject alcohol
into body cavity, or eviscerate specimen. Never use
formalin to preserve skins or skeletal material.

a. Stomach contents: Examine and weigh stomach contents. If closer examination is needed, store contents
in 70% alcohol or methylated spirits. Remember tolabel. Collect any endoparasites in gut, and store in
70% alcohol or 10% buffered formalin.

d. Skeletal material: Save at least the skull and if possible
all of the skeleton. Disarticulate limbs and skull from
specimen and remove soft tissues and rub salt over surfaces. Remove brain with forceps. Let bones dry in

315Appendix

2. Scien tit% Postmortem

tographs as described above. After tranquilizing
the cat, take 5- 10 cm3 of blood by superficial
vasopuncture with an appropriate needle orsyringe. Blood should be immediately passed
slowly into vacutained glass tubes containing
EDTA (purple top) and mixed for 2 to 5 minutes toprevent clotting.

b. Other internal parasites: These should be noted andcounted at each site within the body. Representative
samples should be taken and preserved in 70% alcohol.
c. Intestinal length: For FeZis silvestris, to aid determination of hybridization, measure intestinal length frompyloric to anal sphincter in a straight line (see European
wildcat species account).
ii.
d. Other internal parasites: Store uterus and gonads in
10% buffered formalin, 70% alcohol, or methylated
spirits.
.. .

4. Genetic

111.

Three to eight of these tubes should be centrifugedat 1,500 rpm or greater for 10 minutes. The top
layer of plasma can be removed from each tube
with a plastic pipette and placed in a fresh labelled
Eppendorf tube. These four tubes (two of plasma,
two of cell pellets) should be sealed and stored
frozen. This last step can be delayed for up to 48
hours, if the Eppendorf tubes are kept cool (notfrozen).

iv.

For field biologists without refrigeration, 5-10 mlof whole blood can be placed in equal volume of
the following preservative solution: 100 mm tris
pH. 8.0, 100 mm EDTA, 2% SDS (Sodium
Dodecyl Sulfate), mixed thoroughly. This is theleast desirable form of preservation.

V.

The whole blood tubes (see step iii) can be used for
DNA analysis and the remaining tubes for virus
serology, biochemical analysis, etc.

samplesa. Tissue samples from dead cats:
1.

ii.

Place half-dollar size piece (50 g, l-2 cm diameter)of heart, tongue, skeletal muscle, kidney, liver (inorder of preference) or any other tissue in a ziplock plastic bag. Label and freeze as soon as possible. If you want to take only one sample, heart or
skeletal muscle is best. For liquid nitrogen storage wrap samples in foil or place in cryo-safe
freeze vials.

If there is no access to refrigeration, chop up samples into 1 mm pieces and place in a container withpreservative. The following formula is preferred:
100 mm tris pH. 8.0, 100 mm EDTA, 2% SDS(Sodium Dodecyl Sulfate), mixed thoroughly.
This formula can be obtained from R. Wayne at theaddress below, who will send solution or dry
reagents measured for 500 ml volume of DH20.
These reagents are also available from most university laboratories.
If this preservative is notavailable, 90% alcohol can be used, although it is
not as good.

Blood should be withdrawn from a glass tube witha plastic pipette, and approximately 1 cm3 placed
into 5- 10 separate plastic Eppendorf tubes. Eachtube should be carefully labelled.

The authors’
specimens.

institutions

will gladly accept biologicalAndrew C. Kitchener, Dept. of Natural History, Royal
Museum ofScotland, Chambers Street, Edinburgh EHl
IJF, U.K.

Remember: any tissue that was once living can be a sourceof DNA. New techniques allow geneticists to obtain
potentially useful material from bone, skin, hair, feces,even if the material is several years old and decayed.
Don’t throw anything away if it may be important!

Steven McOrist, Dept. of Veterinary Pathology, Universityof Edinburgh, Veterinary Field Station, Easter Bush,
Midlothian EH25 9RG, U.K.

b. Blood samples from live cats:i.

Robert K. Wayne, Nuffield Laboratories,
Institute ofZoology, Zoological Society of London, Regent’s Park,
London NW1 4RY, U.K. U.S. address:Dept. of Biology,
621 Circle Drive South, University of California at Los
Angeles, LosAngeles, CA 90024.

If you are routinely trapping cats for radio collaring or tagging, it is possible to take blood samples
for DNA and other blood analyses. Remember torecord measurements, weight, and take pho-316

Appendix

AppendixCommonName

Addax
AgoutiAndean fox
Arabian red-legged partridge
ArgaliArgus pheasantArmadillo
Asiatic ibex
Asiatic red sheep
Asiatic wild ass/kiang
Bamboo rat
Banded crake
Big-eared climbing rat
Bighorn sheep
BlackbuckBlack koorhanBlack rat
Blue sheep/bharal
Brown-earedbulbulBuff alo
Camel
Cane mice
Cape fur seal
Cape hare
Capuchin monkeys
Capybara
Caribou, reindeercw
ChamoisCharacid fish
Chevrotains
Chinkara
ChitalChukor partridge
Common duiker
Cotton rats
Cottontail rabbits
Coypu
Dall sheep
Dik-dik
Dorcas gazelle
Dung beetles
Eland

Scientificorder by common

CommonName

Addax nasomaculatus
Dasyprocta variega ta
Dusicyon culpaeus
Alectorus melanocephalaOvis ammon
Argusianus argus
Dasypodidae
Capra ibex sibirica
Ovis orien talis
Equus kiang
Rhizomys sinense
Porzana fusca
Oto tylomys phyllo tis
Ovis canadensis
Antilope cervicapra
Eupodotis afra
Rattus rattus
Pseudois nayaur
Hypsipetes amaurotis
Syncerus caffer
Camelus dromedarius
Zygodontomys
Arctocephalus pusillus
Lepus capensis
Cebus spp.
Hydrochaeris hydrochaeris
Rangifer tarandus
Galea spixi
Rupicapra rupicapra
Characidae
Tragulidae
Gaze/la bene tti
Axis axis
Alectorus chukor
Sylvicapra grimmia
Sigmodon spp.
Sylvivagus spp.
Myocastor coypus
Ovis da/Ii
Madoqua kirki
Gaze/la dorcas
Scarabaeinae
Tauro tragus oryx

of Species

in Text

in the Text

name)

Name

Elephant, African
Elephant, Asian
Elk/Wapiti
European hare
European rabbit
Fallow deer
Gaur, seladang
GemsbokGerbils
Gerenuk
Giant panda
Gibbons
Goral
Great gerbil
Grey fox
Grey jungle fowl
GuanacoGuinea pigs
Guinea fowl
HamsterHartebeest
Harvester termite
Hog deer
Ibex
ImpalaIriomote dwarf hog
JerboasKangaroo rats
Kishinone skink
Kob
Kudu, greater
Kudu, lesser
Land tortoises
Lesser anteaters
MarkhorMarmosets
Marmots
Marsh rat
Mole rats
Moose
Mouflon
Mountain beaver
Mountain chinchilla

317

Names

3

Scientific Names of Species Mentioned(in alphabetical

3. Scientific

Scientific

Name

Loxodonta africana
Elephas maximus
Cervus elaphas
Lepus europaeus
Oryctolagus cuniculus
Dama damaBos gaurus
Oryx gaze/laGerbillus/Meriones
spp.
Litocranius walleri
Ailuropoda melanoleuca
Hylobates spp.
Nemorhaedus goral
Rhombomys opimus
Dusicyon griseus
Gallus sonnerati
Lama guanicoeCavia spp.
Numida meleagris
Phodopus spp.
Alcelaphus spp.
Hodotermes mossambicus
Axis porcinus
Capra ibex
Aepyceros melampus
Sus scrofa riukinanas
Dipodidae
Dipodomys spp.
Eumecas kiskinovi
Kobus kobTragelaphus strepsiceros
Tragelaphus imberbis
Geochelone spp.
Tamuanda spp.
Capra falconeri
Callitrichidae
Marmota spp.
Holichilus brasiliensi
Myospalax spp.
Alces alces
Ovis orien talis
Aplodon tia rufa
Chinchilla bre vicauda ta

Appendix

Common3. Scientific

Namesof Species

Name

Mountain reedbuck
Mountain viscachas
Mouse deer
Mouse opossum
Muntjac/barking
deer
Musk deer
Nile grass rat
NilgaiOpossum
Oribi
Pangolin, African
oryx
Paca
Peccaries
Penguin (Patagonia spp.)
Pig-tailed macaque
Pikas
Porcupine, Indian
Porcupine, North American
Porcupine, prehensile-tailed
Proboscis monkey
Pronghorn
Puku
Red bracket deer
Red deer
Red duiker
Red hartebeest
Reindeer, caribou
Rhino, black
Rhino, white
Rice rat
River turtle
Roan antelopein Text

Scientific

Name

CommonRedunca fulvorufula
Lagidium spp.
Tragulus spp.
Marmosa spp.
Muntiacus spp.
Moscus spp.
Arvican this nilo ticus
Boselaphus tragocamelus
Didelphidae
Orebia ourebi
Manis spp.
Oryx leucoryx
Agouti paca
Tayassu spp.
Eudyptes spp./Spheniscus
magellanicus
Macaca nemestrima
Ochotona spp.
Hys trix indica
Erethizon dorsatum
Coendou prehensilis
Nasalis lavatus
An tilocapra americana
Kobus vardoni
Mazama americana
Cervus elaphas
Cephalophus nigrifrons
Alcelaphus buselaphus
Rangifer tarandus
Diceros bicornis
Cera to therium simum
Oryzomys spp.
Podocnemis unifilis
Hippotragus equinus

NameRock hyraxes
Roe deer
Ryuku flying fox
Sable antelope
SambarSand fox
Snowcocks
Snowshoe hare
Spiny pocket rats
Spiny rats
Springbok
Spring hare
Squirrels
Squirrel monkeys
Striped hyaena
Susliks
Tapir
Thomson’s gazelle
Three-toed sloth
Tolai hare
Tree hyrax
Tsama melon
Tufted deer
Vlei (swamp) rats
Warthog
WaterbuckWhite-tailed deerWhite-tailed pine voleWild pig
Wildebeest
Wolf
Wood rats
Yak
Zebra

318

Scientific

Name

Procaviidae spp.
Cervus capreolus
Pteropus dasymallus
Hippotragus niger
Cervus unicolor
Vulpes riippelli
Tetraegallus spp.
Lepus americanus
Heteromys spp.
Proechimys spp.
Antidorcas marsupialis
Pedetes capensis
Sciurus spp.
Saimiri spp.
Hyaena hyaena
Spermophilus spp.
Tapirus spp.
Gaze/la thomsoni
Bradypus tridactylus
Lepus capensis tolai
Dendrohyrax
Citrullus lana tus
Elaphodus cephalophus
Otomys spp.
Phacochoerus
aethiopicus
Kobus ellipsiprymnus
Odocoileus virginianus
Pitymys leucurusSus scrofa
Connochaetes
spp.
Canis lupus
Neotoma spp.
Bos grunniens
Equus spp.

Appendix

Appendix
Species-Habitat

I. Species

Association

Habitat

Associations

4

Associations

This Appendix is divided into four sections. The first section lists the species associated with each habitat type. The
second section contains the same information organizeddifferently: habitat associations by species. Section 3
describes the methodology for calculating species range
size (for vulnerability ranking), while the fourth section
describes the methodology for estimating percentage of
species range that is protected.

This Appendix contains the species-habitat associations
which were used to rank cat species vulnerability andassess global habitat status and protected area coveragewith respect to cat conservation (Part II, Chapter 1). The
Habitat chapter describes how species associations wereassigned. When reading through this Appendix, reference
should also be made to Figures 2a-f and Tables l-2,4, 8,
and 11 in the Habitat chapter. These figures map the global
habitat types of Olson et al. (1983); provide a definition ofeach type; list the number of species associated with each
type; indicate rate of change in the geographic area of eachtype since 1980; and list percentage of range area protected
for each cat species.

Section4. Species-

Readers should correspond with the Chairman of the Cat
Specialist Group to correct any errors in species-habitat
association, and thus improve its application as a conservation tool.

by Habitat TypeSpecies codes are used rather than full scientific names. The code consists of the first three letters of thespecific name (e.g., par = Panthera pardus, leopard). Two other species names also begin with these letters, so theircodes are: pdl (Leopardus pardalis, ocelot) and pdn Lynx pardinus, Iberian lynx). Similarly, tig is Panthera tigris (tiger)
and tgn Leopardus tigrinus (oncilla); mar is Felis margarita (sand cat) and mma Pardofilis marmorata (marbled cat).
Species are listed by species code in alphabetic, rather than generic, order.

Degree of Species Association by Habitat Type
StrongMajor forest and woodland (1 and 2)
aur, bad, ben, can, car, cha, con,geo, gui, jub, leo, lyn, mma, neb,
one, par, pdl, pdn, pla, rub, ruf,
ser, sil, tern, tig, tgn, viv, wie, yag.
1. Closed forest and woodlandaur, bad, ben, can, con, gui, leo,lyn, mma, neb, one, par, pdl, pla,
ser, tern, tig, viv, wie.
Taiga and other conifer
can, con, lyn.

Significant

Marginalcol.

bie, uric.

car, cha, geo, rub, ruf,
sil, tgn, yag.

bie, col, jub, pdn, uric.

ben, par, ruf, tig.

bie, pdn, sil, tern, uric.

319

Appendix

4. Species-Habitat

Associations

Marginal

Significant

Strongla. Main and southern taigacan, lyn.

uric.

1b. Other conifer
can, con, lyn.

ben, par, ruf, tig.

bie, pdn, sil, tern, uric.

con, geo, neb, par, ruf,,
ser, sil, tern, tig.

can, car, cha, col, pdl, pdn,
uric, yag.

ben, geo, lyn, par, ruf, sil.

cha, col, con, neb, tern, tig.

con, neb, par, ruf, ser,tern, tigcan, car, cha, col, pdl, pdn,
sil, uric, yag.

Main tropical/subtropicalbroad-leaved forest
aur, bad, ben, cha, con, leo, mma,
neb, one, par, pdl, pla, ser, tern,tig, viv, wie

car, rub, sil, tgn, yag.

jub.

1e. Broad-leaved humid forest
aur, bad, ben, con, mma, neb, one,par, pdl, pla, tig, tern, viv, wie.

cha, tgn, yag.

Mid-latitude broad-leaved
and mixed forest
ben, gui, lyn.

1c. Temperate broad-leaved forest

Id. Mixed woods
ben, gui, lyn.

15 Tropical dry forest and woodlandcha, leo, par, ser, tern, tig, viv.

2. Open or interrupted woodlandcar, geo, jub, leo, one, par, pdl,
pdn, rub, ruf, ser, sil, tgn, yag.

ben, car, con, mma, neb,
pdl, rub, sil, yag.

jub, one, tgn, wie.

aur, bad, ben, can, cha, col, con,gui, lyn, mma, neb, pla, tern, tig,viv, wie.

2a. Tropical savanna and woodlandcar, leo, one, par, pdl, ser, sil, yag.

aur, con, jub, rub, wie.

cha, col, geo, ruf, viv.

aur, bad, neb, ser, tern, tig.

ben, car, cha, col, con, mma,
par, pdl, wie, yag.

cha, col, con, gui, leo, one,par, pdl, ruf, sil.

ben, ser.

2b. Tropical montane complexes

tgn
2c. Other dry woods/scrub/grasscar, geo, jub, pdn, rub, yag.

complexes

2d. Northern or maritime taigacan, lyn.

320

Appendix

Strong2e. Second growth woods andfield mosaics
ruf, sil.

4. Species-Habitat

Associations

Significant

Marginalben, can, cha, col, geo, lyn,rnma, rub, par, pla, ser, tern,
tgn, viv, yag.

aur, car, con, jub, leo, neb, one,pdl, pdn, tig, wie.

ben, cha, geo, rub, ser, viv.

can, car, col, con, gui, jub, lyn,ruf, par, sil, yag.

ben, cha, rub, viv.

par.

cha, geo, rub, ser.

ben, can, car, col, con, gui, jub,lyn, ruf, par, sil, yag.

jac, leo, pdl, rub.

ben, can, gui, lyn, mar, neb, one,
ser, tern, tig, uric.

jac, leo, pdl, rub.

ben, gui, lyn, mar, neb, one, ser,tern, tig, uric.

3. Settled areas: cropped, residential,commercial and associated marginal lands

3a. Paddyland

3b. Towns, farms and otherirrigated dryland row crops

4. Grass and shrub complexes:
low vegetation with few or no trees
bie, car, cha, col, con, geo, jub,man, nig, par, pdn, ruf, sil, yag.
4a. Grassland or shrublandbie, car, cha, col, con, geo, jub,man, nig, par, pdn, ruf, sil, yag.
4b. Cold grass or stunted woody complex

bie, can, lyn, uric.
5. Tundra, desert and semi-desert
car, col, geo, jac, man, mar, nig,sil, uric.

bie, con, jub, leo, par, ruf.

5a. Tundra
jac, uric.

*.

can, cha, lyn.

bie.

can, col, con, lyn, man, par.

Desert and semi-desert
car, col, geo, man, mar, nig, sil.

con, jub, leo, par, ruf.

bie, cha, lyn, uric.

5c. Cool semi-desert scrubcol, geo, man.

con.

lyn, ruf, sil, uric.

5d. Sand desert
mar.

car, nig, sil.

jub, leo, man, par.

Se. Other desert and semi-desert
car, mar, nig, sil.

jub, leo, man, par, ruf.

bie, cha, con, lyn.

6. Major wetlandscha, one, pla, ser, tig, viv.

pdn.

aur, ben, con, neb, pdl, ruf, yag.

321

Appendix

4. Species-Habitat

Associations

Section 2. Habitat Association

by Species

Habitat types are abbreviated by alphanumeric code: see brevious section for kev.
Taxa

Degree of Habitat Association
Significant
Marginal

Stronglf, 2e, 3b, 5d

Sub-Saharan Africa
N. Africa-SW Asia
Global

2c, 4a4a, 5e2c, 4a2a, 5e2c, 5d2a, Self, 2e, 3b, 5d

Sub-Saharan Africa
N. Africa-SW Asia
Global2a, 2c, 4a
4a, 5e2a, 2c, 4a, 5elf, 5e2a, 2c, 5d
lf, 5d2b, 2e, 3b, 5d
Id, 2e, 3b
Id, 2b, 2e, 3b

C. badiale2b

C. temminckile, IfId, 2b, 2elb, lc, 4a

F. bieti

4a5alb, 4b, 5e

4a, 6
If, 6
lf, 4a, 6

2c, 2e, 3b
le, 2e, 3a, 3b
le, 2c, 2e, 3a, 3blc, Id, 5elc, Id, 2a, 2b, 4alc, Id, 2a, 2b, 5e

A. jubatus

C. caracal

F. chaus

N. Africa-SW Asia
Tropical Asia
Global4a

F. margarita

5d, 5eF. nigripes

4a, 5e5d

2a, 4a, 5e
4a, 5elc, 2e2a, 2e, 4a, 5elf, 2c, 5d5d2c, 4alc, lf, 2c, 5d

Id, 2e, 3b2c, 2e, 3b, 5c
lb, 3blb, Id, 3b, 5c

H. yaguarondi

2a, 2c, 4ale, lf, 2e

Id, 2b, 3b, 6

L. pardalisle, 2alf, 2c, 4a

Id, 2b, 2e, 6

L. tigrinus

2ble, 2elf

L. wiedii

le2alf, 2b, 2elf, 2a, 6
Id
If, 2a, 6

2b, 2e, 3b2c, 4aId, 2b, 2e, 3b2c, 4ala, lb2d, 2eId, 3b, 4b, 5ala, lb, Idla, lb, Idla, lb, Id

lc, 2d, 2elc, 2elc, 2d, 2e

4a, 4b, 5a, 5c, 5e3b3b, 4a, 4b, 5a, 5c, 5e

2c, 4a6lb, Id, 2e

F. silvestris

L. serval

lybicaornatasilvestris
GlobalSub-Saharan Africa
N. Africa-SW Asia
GlobalL. canadensis
L. lynx

L. pardinus

Asia
EuropeGlobal322

2c, 4aAppendix

Degree of Habitat Association
Significant

Taxa
Strong4. Species-

Habitat

Associations

Marginal

L. rufus

2e, 4alb, lc, Id, 2c, 5e

2a, 3b, 5c, 6

0. colocolo

4a, 5c2c, 2elc, Id, 2a, 2b, 3b, 5a

0. guigna

Id

2c3b, 4a0. geoffroyi

2c, 4a, 5c

lc, 2e, 3b2a0. jacobitus

5a4a0. manul

4a, 4b, 5c

5e

5a, 5dP. bengalensis

Id, lelb, lc, If, 2e, 3a2b, 2c, 3b, 4a, 6

P. planiceps

le, 6

2e

P. rubiginosus

2clf, 2a, 2e, 3a, 3b, 4a

P. viverrinus

le, lf, 6

2e, 3a2aP. aurata

le2a, 2b2e, 6

P. concolor

lb, le, 4a

Id, 1f, 2a, 2c, 5clc, 2b, 2e, 3b, 5a, 5e, 6N. nebulosa

leId, lf, 2blc, 2e, 4a, 6

P. leoIf, 2a2c, 4a, 5e

2e, 5dP. oncale, 2a, 62clf, 2e, 4ale, lf, 2a4ale, Ifle, lf, 2a, 4a2c, 2e, 4a, 5elc, 2a, 2c, 5elb, Id, 2b, 2elb, lc, Id, 2c, 2e,2b, 3b, 5d
Id, 2e, 5dlc, 2a, 3a, 3b, 5a2b, 3a, 3b, 5a, 5d, 5e

P. tigrisle, If, 6

lb, Id, 2blc, 2e, 4a

P. marmorataleIf, 2e2bU. uncia

5a

P. pardus

Sub-Saharan Africa
N. Africa-SW Asia
Tropical Asia
Globalla, lb, Id, 4a, 4b, 5c

323

Appendix

4. Species-

Habitat

Associations

33% to eliminate the Sahel region. There is a high potential for error in applying reduction factors, as they were
calculated by visual comparison of species range maps tothe habitat types, taking into account patchy distribution or
rarity as described in the species accounts.
Only one other source was used for data on habitat area,
and then only for a few species. WCMC (1992) combines
tropical forest area for Java, Sumatra, and Borneo withdata for other Indonesian islands and Peninsular Malaysia.
When separate habitat area data was required, figures fromthe KJCN Conservation Atlas of Tropical Forests: Asiaand the Pacific (Collins et al. 1991) were used.
In the few cases where good data available on the areaof species contemporary range (snow leopard [Fox 19951
and Iberian lynx [Rodriguez and Delibes 1992]), ourrange area estimates have turned out to be too high. The
overestimate was by a factor of 0.25 in the case of thesnow leopard, and by 5.3 in the case of the Iberian lynx.
Error in the case of the Iberian lynx is unavoidable as themethodology used here lacks resolution on a small scale.
The actual range of the Iberian lynx is estimated at only
14,000 km2, and the lynx is restricted largely to the southof the Iberian peninsula in over 100 isolated sub-populations. Nonetheless, the estimate served its purpose by
qualifying the geographic range of the Iberian lynx for thesmallest category, “Restricted.”
The purpose of the exercise was to compare species
range on a relative, rather than absolute scale. While it ishoped that the bias for other species is closer to that of thesnow leopard range calculation, this cannot be determinedwith the data at hand. The error arises from many sources,
including: (1) the methodology of Olson et al. (1983),
which assigns a 3,050 km2 pixel to the habitat type predominating within it, regardless of the other types of habitat it contains; (2) errors in geographic area calculations
which may have arisen when WCMC (1992) compiled
their table; (3) errors in assigning species-habitat associations and degree of association; (4) errors in applicationof reduction factors; and (5) failure of the species range
maps to represent actual species range. Given the likelihood of error, our estimates of species range area should
not be treated as definitive. They represent a first attempt
to apply a standard methodology to estimation of cat
species range, and are useful for developing a more strategic approach to cat conservation. It is hoped that they
serve as a stimulus for further work.

Section 3. Methodology
Used toCalculate Species Range Area
Approximate range sizes of the cat species were presented
in Box 1 in the Introduction to the Species Accounts (PartI). Species range area was calculated by adding up thegeographic coverage of all habitat types within eachspecies’ contemporary range with which the species isstrongly or significantly (not marginally) associated.
Figures for geographic coverage by habitat type wereobtained from two sources. The primary source wasTable 18.2 in WCMC’s Global Biodiversity (WCMC
1992), which combines the habitat types of Olson et al.
(1983) into eight major groupings (Major forest,
Interrupted woods, Crop and settlements, Grass andshrub, Polar and alpine, Desert and semi-desert, Major
wetlands, and Other coastal aquatic). The habitat type
“Other coastal aquatic” was not used for species-habitat
association or species range area calculation because it
includes the area taken up by lakes and beaches. The
WCMC table lists percentage area occupied by these eight
habitat groupings for each country (Saudi Arabia, forexample, is listed as 62% Desert and semi-desert). The
percentage area was converted to km2 using data on country area given in The Times Atlas of the World (8th edn.,1990). Area was expressed in millions of km2, rounded
off to three decimal places (e.g., Saudi Arabia, with a total
land area of 2.4 million km2, has 1.490 million km2 ofdesert and semi-desert).
In many cases, a reduction factor was applied toaccount for either: (1) species occurrence in only a portion of a country’s habitat type, or; (2) species occurrence
in only a portion of WCMC’s (1992) habitat type groupings. Examples of the first type include the tiger’s occurrence in only a tiny portion of Russia’s coniferous forest(1 b), and the sand cat’s apparent absence from much of
Egypt. An example of the second type is the leopard’s
scattered, patchy distribution in the sub-Saharan African
Sahel region (a portion of habitat type 2~). These three
examples also illustrate the three ways in which reduction
factors were applied. In the case of the tiger, pixels ofconiferous forest were counted on the habitat map andmultiplied by 3,000 km? In the second case, the sand cat,
the geographic area of desert and semi-desert for Egypt
was reduced by 80%. In the leopard’s case, the total areaof habitat type 2c for sub-Saharan Africa was reduced by324

Appendix

4. Species-Habitat

Associations

tected areas falling within a species range contained thespecies. The approximate total number of protected areas
which potentially contain cat species, and the approximate total geographic area covered by these protectedareas, were obtained by adding up the numbers for eachbiogeographic province falling within a species range. If
the range of a particular cat species included only a portionof an Udvardy biogeographic province, a corresponding
reduction factor was applied to both the number and total
area of the protected areas of the biogeographic province.
The percentage of protected species range was obtained by
dividing the total area protected by the total estimatedspecies range size.

Section 4. Methodology
Used toCalculate Species OccurrenceinProtected Areas
Species occurrence in protected areas is presented in Part
II, Chapter 1, Table 11. WCMC (1992: Table 29.5) calculated the number and total geographic area of protectedareas occurring within the biogeographic provinces of theworld, as defined by Udvardy (1975). Udvardy (1975)
delineated 195 biogeographic provinces, which do not correspond to the habitat types defined by Olson et al. (1983).
However, from Udvardy’s (1975) map, it is possible toidentify the biogeographic provinces which fall within cat
species ranges. The assumption was made that all pro-325

Appendix

5. Cat Specialist

Group

Members

Appendix

5

Cat Specialist Group Members
Chair: Peter Jackson; Deputy
Chair: Alan Shoemaker;
Vice-Chair(Asia): Valmik Thapar; Vice-Chairs(Cheetah):
Laurie and Daniel Marker-Kraus;
Vice-Chairs(Lynx):Urs and Christine Breitenmoser;
Vice-Chair(Projects):
Kristin Nowell;Vice-Chair(Snow leopard):
Helen Freeman;
Research
Librarian:
Gail Foreman

Agadjanian, Frunze S.
Institute of Zoology, Armenian
Academy of Sciences, P. Sevak
Str. 7, Erevan 375 044, Armenia
Ahlborn, Gary
Biosystems Analysis, 17 145 Park
Avenue, Sonoma, CA 954768506
U.S.A.
Ahmed, Ashiq
WWF Pakistan, Univ. of Peshawar,
UP0 Box 1439, Peshawar, Pakistan
Alwis, Lyn de
30 Hotel Road, Mount Lavinia,
Colombo, Sri Lanka

Bertram, Brian
Fieldhead, Amberley, Stroud,GL5 5AG, GB
Biquand, Sylvain
42 r. des Boulangers, Paris 75005,
FranceBlomqvist, Leif
Helsinki Zoo, Korkeeasaari,
Helsinki SF 005706, Finland
Borner, Markus
Regional Representative, Frankfurt
Zoological Society, P.O. Box 3 134,
Arusha, Tanzania

Amrasanaa, G.
Mongolian Academy of Sciences,
Ulan Bator 5 1, Mongolia

Bothma, J. du P.
Centre for Wildlife Research,
University of Pretoria, Pretoria
0002, South Africa

Aranda, Marcel0
Instituto de Ecologia, Apdo. Postal
63, Xalapa 9 1000, Mexico

Bragin, Anatoley P.
Trudovoye, K.Tstetkin 38,
Vladivostok 692 806, Russia

Artois, Marc
CNEVA, B.P. 9, Malzeville 54220,
FranceBreitenmoser, Urs
Villettengassli 4, Muri 3074,
Switzerland

Bailey, Theodore N.
Kenia National Wildlife Refuge,
POB 2139, Soldotni 99669,
Alaska, U.S.A.

Brockelman, Warren Y.
Faculty of Science, Mahidol
University, Rama VI Road,
Bangkok 10400, Thailand

Belden, Robert
Florida Panther Recovery Unit,
Florida Game and Fish Comm.,
4005 South Main Street,
Gainesville, FL 32601, U.S.A.

Brooks, Daniel M.
Houston Zoological Gardens,
15 13 N. MacGregor, Houston,
TX 77030, U.S.A.

Berry, H.H.
Nature Conservation Directorate,
Namib Research Institute, P.O. Box1204, Walvis Bay 9190, Namibia

Capt, Simon
Gheiweg 55, Einigen 3646,
Switzerland

326

Caro, Tim
Wildlife and FisheriesDept,
University of California, Davis,
CA 95616, U.S.A.
Chakrabarti, Kalyan
ConservatorForests,Research,
Bikash Bhavan, North Block,3rd Floor, Calcutta 700 091, India
Chavda, Divyabhanusinh
Vice-President, Northern Region,
Taj Group of Hotels,
1 Man Singh Road,New Delhi
110 011, India
Chellam, Ravi
Wildlife Institute of India,
P.O. Box 18, Chandrabani,
Dehra Dun 248 001, India
Chundawat, Raghunandan
Wildlife Institute of India,
P.O. Box 18, Chandrabani,
Dehra Dun 248 001, India
Cop, Janez
Verovskova 43, Ljubljana, Slovenia
Crawshaw, Peter G.
C.P. 750, PR 85890,
Foz do IguaCu,Brazil
Dao Van Tien
Laboratory of Zoology, 19Le
Thanh Tong Street, P.215 Khu
Thanh Cong, Hanoi, Vietnam
Davidson, Bruce
Dept. of Medical Biochemistry,
Medical School, University of
Witwatersrand, Johannesburg2193,
South Africa

Appendix

Delibes, Miguel
Estacion Biologica de Donana,
Apt. 1056 Pabellon de1 Peru,
Avenida de Maria Luisa s/n,
Seville 4 1080, Spain

Freeman, Helen
President, Intemat. Snow Leopard
Trust, 4649 Sunnyside Av. No.,
Seattle, WA 98103, U.S.A.

Dragesco- Joffe, Alain
Villa La Rousseraie, Chapellede Rousse, Juraqon, France

Garcia-Perea, Rosa
Museo National de Ciencias Nat.,c/ J. Gutierrez Abascal,
2, Madrid 28006, Spain

Dunishenko, Yuri
Wildlife Management Inst.,
P.O. Box 1769, L. Tolstoy St. 15-A
Khabarovsk 680 049, Russia

Ghosh, Arin
Director, Project Tiger, Bikaner
House Annexe 5, Shah Jahan Road,New Delhi 110 011, India

Dunstone, Nigel
Department of Zoology, Universityof Durham, South Road, Durham
DHl3LE, GB

Gogate, M.G.
Conservator of Forests, Research
Circle, New PMT Building,
3rd Floor, Pune 411 042, India

Eisenberg, John F.
Dept. of Natural Sciences,
The Florida State Museum,
University of Florida, Gainesville,
FL 32611, U.S.A.

Grisham, Jack
General Curator, Oklahoma
Zoological Park, 2 10 1 NE 5Oth,
Oklahoma City, OK 73 111, U.S.A.

Foose, Tom
Int. Rhino Foundation, c/o The
Wilds, 85E Gay St., Suite 603,
Columbus, OH 43215, U.S.A.
Foreman, Gail
Utica Zoo, Steele Hill Road,
Utica, NY 13501, U.S.A.
Fouraker, Michael D.
Asiatic Lion Studbook Keeper,
Fort Worth Zoo, 1989 Colonial
Parkway, Fort Worth, TX 76110,
U.S.A.
Fox, Joseph
Department of Ecology, Universityof Tromso, P.O. Box 3085 Guleng,
Tromso 9001, Norway
Frame, George
P.O. Box 822, Cape May Court
House, NJ 08210, U.S.A.
Franklin, William L.
Dept. of Animal Ecology,
124 Sci II, Iowa State University,
Ames, Iowa 50011, U.S.A.

Gui Xiaojie
Forestry Department, Hunan
Province, 39 Nan Da Road,
Changsha, China

5. Cat Specialist

Group

Members

Hornocker, Maurice
Homocker Wildlife ResearchInstitute, P.O. Box 3246, University
Stn, Moscow, Idaho 83843-0246,
U.S.A.
Ilany, Giora
Sapir 86825,Israel
Izawa, Masako
Dept. of Biology, Faculty of
Science,University of the Ryukus,
Nishihara-oho,Okinawa 903-O1,
Japan
Jackson, Peter
Route desMacherettes,
Bougy-Villars 1172,SwitzerlandJackson, Rodney
18030ComstockAvenue,
Sonoma,CA 95476,U.S.A.
Jalkotzy, Martin
ARC Assoc. ResourceConsultants,
2201 34th Street SW, Calgary
T3E 2W2, Canada
Jenny, David
Rechengasse
2,562O Bremgarten,
Switzerland

Ha Dinh DueCentre Natural Resources Manage,
University of Hanoi,
19 Le Thanh Tong St, Hanoi,
VietnamJewell, Peter
South Barn High Street, Fen Ditton,
Cambridge,CB3 8ST, GB

Halloy, Stephan R.P.
Crop and Food Research, Invermay
Agricultural Centre, Private Bag
50034, Mosgiel, New ZealandJohnsingh, A. J.T.
Wildlife Institute of India,
P.O. Box 18, Chandrabani,
Dehra Dun 248 001, India

Hemmer, Helmut
Anemonenweg 18, MainzEbersheim 6500, Germany

Johnson, Kurt A.
1312Oak Ridge Ave., Apt 112,
EastLancing, MI 48823, U.S.A.

Herrenschmidt,
Veronique
Ministere de L’Environnement,
Dir. Protection de la Nature,
14 Bd General Leclerc, Neuilly
92524, France

Johnson, Warren E.
Lab. of Viral Carcinogensis,
National CancerInstitute, Building
560, Frederick, MD 21701-1201,
U.S.A.

Hoogesteijn, Rafael
Apartado 3083, El Trigal, Valencia,
Edo. Carabobo, Venezuela

Kaczensky, Petra
Fichtenstr. 39, Putzbrunn 8011,
Germany327

Appendix

5. Cat Specialist

Group

Members

Karanth, K. Ullas
Associate Research Zoologist,
Wildlife Conservation Sot. (NYZS),
499 Chitrabhanu Rd, Kuvempu
Naga, Mysore 570 023, India
Khan, Anisuzzaman
13 Monipuripara, Serebangla
Nagar, Dhaka, Bangladesh
Khan, M.A. Reza
Director, Dubai Zoo,
P.O. Box 67, Dubai, Dubai (UAE)
Khan, Mohammed
Ibu Pajabat Jabatan Perhutanan,
5th Floor, Wisma Sumbar Alam,Jalan Stadium Petra Jaya, Kuching
93600, Sarawak, Malaysia
Kitchener, Andrew
Royal Museum of Scotland,
Chambers Street, Edinburgh EH 1
GB

Kvam, Tor
Norsk Institutt, for Naturforskning,
Tungasletta 2, Trondheim 7004,
NorwayLaurenson, Karen
Upland Research Group,
The Game Conservancy,
Crubenmore Lodge,
Newtonmore PH20 IBE, GB
Lawton, Peter
19 Kautilya Marg, New Delhi
110 021, India
Lewis, John
Director, John Ball Zoological
Gardens, 1300 West Fulton Street,
Grand Rapids, MI 49504, U.S.A.
Leyhausen, Paul
Auf’m Driesch 22, Windeck 5 1570,
Germany

Kock, Richard
Chief Veterinarian, Kenya Wildlife
Service, P.O. Box 40241, Nairobi,
Kenya

Lu Houji
Biological Conservation Centre,
Dept. of Biology, East China
Normal University, Shanghai
200 062, China

Koehler, Gary M.
P.O. Box 1852
Snowqualmie, WA, 98065, U.S.A.

Mallon, David
3, Acre St., Glossop,
Derbyshire SK13 8JS, GB

Korkishko, Viktor G.
Far East Leopard Fund, Kedrovaya
Pad Reserve, St. Primorsky,
khasansky r-on, Primorsky Krai,
Russia 692 710

Marker-Kraus,
LaurieInt. Cheetah Studbook Keeper, P.O.
Box 247, Windhoek 9000, Namibia

Koshkarev, Eugene P.
Tchaikovsky Street 140,
Appartment 6, Irkutsk, Russia
Kotwal, Prem Chand
Associate Professor, Indian Inst.
Forest Management, Nehru Nagar,
P.O. Box 357, Bhopa1462 003,
India
Kumar, Ashok
B5/22 Safdarjang Enclave,
New Delhi 110 029, India

Maruska, Ed.
Executive Director, Cincinnati Zoo,
3400 Vine Street, Cincinnati, Ohio
45220, U.S.A.
Maskey, Tirtha
Director Zoo, Kathmandu, Nepal
Matyushkin, E.N.
Dept. of Biogeography, Geography
Faculty, Moscow State University,
Moscow W-234, Russia 119 899
McDougal, Charles
Tiger Tops, P.O. Box 242,
Kathmandu, Nepal

328

McNeely, Jeffrey
IUCN, Gland 1196, Switzerland
Mellen, Jill
Conservation Research Coordinator
Washington Park Zoo,
4001 SW. Canyon Road,
Portland, OR 9722 l-2799, U.S.A.
Mendelssohn, Heinrich
University of Tel Aviv, Faculty ofLife Sciences, P.O. Box 39040,
Ramat Aviv 69978, Tel Aviv, Israel
Mills, Gus
Kruger National Park, Private Bag
X402, Skukuza 1350, South Africa
Miquelle, Dale
Homocker Wildlife ResearchInstitute, P.O. Box 3246, University
Station, Moscow, ID 83843, U.S.A.
Mishra, Hemanta R.
Environment and Social Affairs
Div, Asia Tech. Dept., The World
Bank, 18 18 H Street NW,
Washington, D.C. 20433, U.S.A.
Miththapala, Sriyanie
176 Polhengoda Road, Colombo 5,
Sri Lanka
Mizutani, Fumi
c/o Lolldaiga Hills Ltd,
P.O. Box 26, Nanyuki, Kenya
Mondolfi, Edgardo
Quinta Masapo, Avenida Norte deAlta Florida, Caracas 1050,
Venezuela
Morsbach, Dieter
Directorate of Resource
Management, Ministry ofEnvironment and Tourism,
Private Bag 13306, Windhoek
9000, Namibia
Mueller, Peter
Zoologische Garten, Pfaffendorger
Str. 29, Leipzig 7010, Germany

Appendix

Myers, Norman
Upper Meadow, Old Road,
Headington, Oxford OX3 8S2, GB
Nader, Iyad A.
Nat. Comm. for Wildlife,
Conservation and Development, PO
Box 61681, Riyadh 11575, Saudi
ArabiaNavarro L, Daniel
Apartado Postal 13, Retorno 8 No.
480A, Fraccionamiento Campestre,
Chetumal, Quintana Roo 77030,
MexicoNorton, Peter M.
Eastern Cape Province, Cape
Nature Conservation, Private Bag
X 1126, Port Elizabeth 6000, South
AfricaNowell, Kristin
2520-4,41st St. NW
Washington DC 20007
U.S.A.
O’Brien, Stephen J.
Laboratory of Viral Carcinogenesis,
National Cancer Research Insti,
Bldg. 560, Room 21-105,
Frederick, MD 21702-1201, U.S.A.

5. Cat Specialist

Group

Members

Peters, Gustav
Zoologisches Forschungsinstitut,
Adenauerallee 1SO- 164, Bonn
53113, Germany

Rathore, Fateh Singh
MAA Forestry Farm, Ranthambhor
Road, PO Sharpur Khilchipur,
Sawai Madhopur, Rajasthan, India

Pikunov, Dimitriy G.
Laboratory of Ecology of Animals,
Pacific Institute of Geography,
7 Radio Street, Vladivostok
690 032, Russia

Rau, Jaime R.
Universidad National, Campus
Omar Dengo, Apdo. 1350-3000,
Heredia, Costa Rica

Poole, Kim G.
Wildlife Management Division,
Renewable Resources,
Govt. Northwest Territories,
Yellowknife, NWT,
Canada XlA 2L9
Quigley, Howard
Homocker Wildlife ResearchInstitute, P.O. Box 3246, University
Station, Moscow, ID 83843-0246,
U.S.A.
Quillen, Pat
Society for Scientific Care, Inc.,
P.O. Box 7535, San Diego, CA
92107, U.S.A.
Rabinovich, Jorge, Dr.
Fundacion Sirena, Casilla 1395,
Correo Central, Buenos Aires 1000,
Argeniina

Rishi, Vinod
Deputy Director, Wildlife Cons.,
Min. Environment and Forests,
Paryavaran, CGO Complex,
Lodi Rd, New Delhi 110 0 11, India
Roelke, Melody E.
Chief Veterinarian, Tanzania
National Parks, P.O. Box 3134,
Arusha, Tanzania
Roy, Deb
24/4 Type 5, Lodi Complex,
New Delhi 110 003, India
Sanyal, Pranabes
57D, Pumadas Road,
Calcutta 700 029, India
Seal, Ulysses S.
Conservation Breeding Specialist
Group, 12 10 1 Johnny Cake Ridge
Road, Apple Valley, MN 55 124,
U.S.A.

Okarma, Henryk
Mammal Research Institute,
Polish Academy of Sciences,
Waszkiewicza 1 D,
Bialowieza 17-230, Poland

Rabinowitz, AlanNYZS/Wildlife Conservation Sot.,
Bronx Zoo, Bronx, NY 10460,
U.S.A.

Ono, Yuiti
Department of Biology, Faculty of
Science, Kyushu University 33,
Fukuoka 8 12, Japan

Ragni, Bernardino
Istituto de Zoologia, Universita
delgi Studi, Via Elce di Sotto,
Perugia 06 100, Italy

Seifert, Siegfried,
Zoologische Garten,
Leipzig 70 10, Germany

Packer, Craig
Dept. of Ecology, Evolution, andBehaviour, 100 Ecology Building,1987 Upper Buford Circle,
St. Paul, MN 55108-6097, U.S.A.

Ranjitsinh, M.K.
Chairman, Narmada Valley Proj,
Narmada Bhavan, Tulsinagar,
Bhopa1462 003, India

Shoemaker, Alan H.
Curator of Mammals, Riverbanks
Zoological Park, P.O. Box 1060,
Columbia, SC 29202- 1060, U.S.A.

Rashid, M. A.
Ketan Appartments, Flat 103,
Fatehganj Camp, Baroda 390 002,
India

Singh, Arjan
Tiger Haven, P.O. Pallia,
Pallia 262 902, India

Panwar, Hemendra S.
Director, Wildlife Institute of India,
P.O. Box 18, Dehra Dun 248 001,
India

329

Seidensticker, John
National Zoological Park,
Smithsonian Institution,
Washington, D.C. 20008, U.S.A.

Appendix

5. Cat Specialist

Group

Members

Slough, BrianFish and Wildlife Branch, Yukon
Renewable Resources, Box 2703,
Whitehorse, YU YlA 2C6, Canada

Tello, Jose Lobao
Technical Director, PDRN,
B.P. 1608, Bangui,
Central African Republic

Smith, James L. David
Dept. of Fisheries and Wildlife,
200 Hodson Hall, 1980 Folwell
Ave, St Paul, MN 55108, U.S.A.

Tewes, Michael
Director, Feline Research Centre,
Texas A&I University, Campus
Box 218, Kingsville, Texas 78363,
U.S.A.

Stahl, Philippe
ONC/Service Technique,
Domaine de Saint-Benoist, 5 rue deSaint-Thibault, Auffargi, Le Perrayen Yvelines 78610, FranceStander, Philip E.
PO Box 17, Grootfontein, Namibia
Stuart, Chris
P.O. Box 6, African Carnivore
Survey, Loxton 6985, South Africa
Sumardja, Effendy
Ministry of Forestry, Bali Prov.,
Complex Niti Mandala, Jalan Raya
Puputan-Renon, Denpasar, Bali,
Indonesia
Sunquist, Melvin E.
118 Newins-Ziegler Hall,
Florida Museum of Natural History.
University of Florida, Gainesville,
FL 32611, U.S.A.
Suvanakorn, Phairot,
Director General, Royal Forestry
Department, Phaholyothin Road,
Bangkhen, Bangkok 10900,
ThailandSwank, Wendell G.
2326 S. Quail Run Road,
Cottonwood, AZ 86326, U.S.A.
Tan Bangjie
Advisor, Chinese Ass. of
Zoological Gardens, 137 Xi Zhi
Men Wai Street, Beijing 100 044,
China
Teer, James G.
Director, R. and B. Welder Wildlife
Fndn, P.O. Drawer 1400, Sinton,
TX 78387, U.S.A.

Thapar, Valmik
19 Kautilya Marg, New Delhi
110 021, India
Tilson, Ronald L.
Director of Conservation,
Minnesota Zoological Garden,
13000 Zoo Boulevard,
Apple Valley, MN 55124, U.S.A.
Tischendorf, Jay
American Ecological Research Ins,
P.O. Box 380, Fort Collins, CO
80522, U.S.A.
Vaughan, Christopher
Posgrado en Manejo de Vida Silv,
Universidad National Heredia,
Campus Omar Dengo,
Heredia 1350, Costa Rica
Villalba, Juan
TRAFFIC Sud America, Carlos
Roxlo 14961301, Montevideo,
Uruguay
Vo QUY
Faculty of Biology, University of
Hanoi, 19 Le Thanh Ton, Hanoi,
VietnamVyas, Mahendra
H-53-D, Saket, New Delhi
110 017, India
Walker, Clive
Lapalala Wilderness Sanctuary,
P.O. Box 645, Bedfordview 2008,
South Africa
Werdelin, Lars
Dept. of Palaeozoology,
Museum of Natural History,
P.O. Box 50007, Stockholm
S-104-05, Sweden
330

Wilson, Vivian J.
Chipangali Wildlife Trust, P.O. Box
1057, Bulawayo, Zimbabwe
Wotschikowsky,
UlrichWildbiologische Gesellschaft,
Postfach 170, D-8103
Oberammergau, Germany
Wright, Anne
Tollygunge Club Ltd, 120
Deshapran Sasmal Road, Calcutta
70003 3, India
Wright, Belinda
Wildlife Protection Society
Thapar House, 124 Janpath,
New Delhi, India 110 001
Xiang Peilun
Municipal Bureau of Parks,
180 Eling Street, Chongqing
630 014, China
Ximenez, Alfred0
Univ. Fed. de Santa Catarina,
Campus Universitario-Trinidade,Caixa Postal 5 132, FlorianopolisSta Catarina 88049, Brazil
Yang Qisen
Institute of Zoology,
19 Zhongguancun Road,
Beijing 100 080, China
Yasuma, Shigeki
c/o PUSREHUT, Univ.
Mulawarman, Gunung Kulua,
Samarinda, Kalimantan-Timur,
Indonesia
Yu Jingping
Dept. of Zoology, Auburn
University, 33 1 Funchess Hall,
Auburn, AL 36849-5414, U.S.A.
Zhyvotchenko, Victor
Science Magazine,
v/o Agroprodmizdat,
Pereckopskaya St, 14 div. 1-133,
Moscow 113 209, Russia
Zuleta, Gustav0
Area Ecologia, Dpto de Biologia,
Univ. de Buenos Aires, Cdad
Universitaria, Pab. II, 4xp,
Buenos Aires 1428, Argentina

Appendix

Appendix
The International

6. IFTF Statement

6

Fur Trade Federation

A Statement

The International Fur Trade Federation (IFTF) represents
national fur trade associations in 28 countries. They
include virtually every country where fur is either seriously produced or traded. Countries in which fur is produced and traded which do not belong to IFTF include
India, Nepal, and most South American countries.

IFTF stands solidly behind practical measures to improvethe enforcement of CITES. It condemns and dissociates
itself from incidents of unscrupulous traders selling threatened species to tourists and indulging in illegal activities.
It is as much in the trade’s interest as in CITES’ intereststhat such practices be stopped.

The Federation’s activities in the field of conservation datefrom the early 1970s when, concerned about the decline
in the flow of leopard skins from producing countries, theIFTF introduced a voluntary ban on the handling of several
cat and other species. Since then, the Federation has
helped to finance a number IUCN and CITES surveys into
the status of various fur-bearing animals. In addition, in
association with the government of Canada, IFTF hashelped to fund important research into improved trapping
methods.

The legitimate international trade has no interest in handling CITES-listed species unless they are available incommercial quantities under proper, regulated controls.
In sum, the trade’s position is straightforward-it
believes
in strictly regulating trade in any species scientifically
proven to be threatened, but expects to trade in the sustainable yield of any species which is not threatened, in
accordance with the sustainable use principles laid down inthe documents, World Conservation Strategy and Caring
for the Earth, published by IUCN-The
World
Conservation
Union, UNEP-the
United Nations
Environment Programme, and MF-World
Wide Fundfor Nature.

In accordance with its 1985 constitution, IFTF is concerned at all times with the conservation of fur-bearing
animals, and supports CITES and the strict observance of
its regulations. IFTF members are bound by this constitution, and may be liable to expulsion if they fail to act
against any member of their own association proven tohave violated CITES regulations.

International Fur Trade Federation
Walton-on-Thames, England

331

Appendix

7. List of Maps,

Figures,and Tables

Appendix

7

List of Maps, Figures, and Tables

Page

Page

Fig. 3. Distribution of the leopard (P. pardus)
in north Africa and southwestAsia ...............................46

Taxonomy of the Felidae
Fig. 1. Branching diagramderived from thephylogenetic tree proposedby Hemmer (1978) ..........xxi

Fig. 4. The undersideof a sandcat’s paw,
showingfur covering ............................................~........47

Fig. 2. Branching diagramredrawn from thecladogramfor extant felids proposedby
Herrington ( 1986)........................................................xxii

Fig. 5. Distribution of the sandcat (F. margarita) ......49
Fig. 6. Distribution of the caracal(C. caracal) in
north Africa and southwestAsia ...................................52

Fig. 3. Phylogenetic relationshipof felid species
basedon a consensusof molecular,karyologic, and ...
morphologicalcharacters...........................................xx111

Part I.

Table 1. Leopard population statusby country.. ......... .45

SpeciesAcccounts

Chapter 3. Tropical Asia

Chapter 1. Sub-Saharan Africa

Fig. 1. Historical distribution of the tiger(P. tigris): mid- 1800s- mid- 1900s...............................59

Fig. 1. Distribution of the black-footedcat (F. nigripes) ...............................................................9

Fig. 2. Presentdistribution of the tiger .........................60

Fig. 2. Distribution of the African
goldencat (P. aurata) .................................................... 11

Fig. 3. Current distribution of the South China
tiger (P. t. amoyensis),with protected areasandproposednetwork ..........................................................62

Fig. 3. Distribution of the cheetah(A. jubatus) in sub-SaharanAfrica ................................ 15

Fig. 4. Distribution of the Amur tiger (P. t. altaica)in the RussianFar East..................................................63

Fig. 4. Distribution of the lion (P. Zeo)in
sub-SaharanAfrica ........................................................20

Fig. 5. Proposedblocks of protected areasfor
tigersin India ................................................................64

Fig. 5. Distribution of the serval (L. serval) ................23

Fig. 6. Possibledistribution of the Bornean
bay cat (C. badia) ....................*..................................... 66

Fig. 6. Distribution and relative abundanceof the leopard (P. pardus) in sub-SaharanAfrica.........27

Fig. 7. Distribution of the cloudedleopard(N. nebulosa).......................................................~.........68

Fig. 7. Distribution of the caracal (C. caracal)
in sub-SaharanAfrica ....................................................31

Fig. 8. Distribution of the Asiatic goldencat(C. temmincki)...............................................................70

Fig. 8. Distribution of the African wildcat(F. silvestris,lybica group) ...........................................34

Fig. 9. Distribution of the flat-headedcat(P. planiceps).................................................................71

Chapter 2. North Africa and Southwest Asia

Fig. 10. Distribution of the rusty-spottedcat(P. rubiginosus).............................................................73

Fig. 1. Pastand presentdistribution of the lion
(P. Zeo)in north Africa and southwestAsia ..................39

Fig. 11. Distribution of the fishing cat(P. viverrinus) ................................................................75

Fig. 2. Pastand presentdistribution of the cheetah(A. jubatus) in north Africa and southwestAsia ...........42

Fig. 12. Distribution of the marbledcat(P. marmorata)..............................................................77332

Appendix

7. List of Maps,

Figures,andTables

Page

Page

Chapter 5. The Americas

Fig. 13. Distribution of the leopard(P. pardus) in tropical and eastAsia .............................80

Fig. 1. Distribution of the kodkod (0. guigna).......... 116

Fig. 14. Distribution of the Amur leopard(P. p. orientalis) in Russiain 1971...............................81

Fig. 2. Distribution of the Andean mountain cat(0. jacobitus) ............................................................... 117

Fig. 15. Distribution of the Amur leopard(P. p. orientalis) in Russiain 1991...............................82

Fig. 3. Pastand presentdistribution and relative
abundanceof the jaguar (P. onca) .............................. 120

Fig. 16. Distribution of the jungle cat (F. chaus)........84

Fig. 4. Possibledistribution of the oncilla(L. tigrinus) .................................................................. 123

Fig. 17. Distribution of the leopardcat(P. bengalensis).............................................................86

Fig. 5. Distribution of the margay (L. wiedi) ............. 125

Fig. 18. Distribution of the Iriomote cat(P. b. iriomotensis)........................................................89

Fig. 6. The IO-year cycle in Canadalynx(L. canadensis)populations........................................ 126
Fig. 7. The relationshipbetweensnowshoehare
and Canadalynx populations...................................... 127

Table 1. Size variation in tiger subspecies...................56
Table 2. Densitiesreported for tigers in
different habitats............................................................61

Fig. 8. Distribution of the Canadalynx(L. canadensis)............................................................ 128
Fig. 9. Distribution of Geoffroy’s cat(0. geoffroyi) .......................................................~.......130

Chapter 4. Eurasia
Fig. 1. Distribution of the snow leopard(U. uncia ....93

Fig. 10. Pastandpresentdistribution of thepuma (P. concolor) ...................................................... 133

Fig. 2. Known distribution of the Chinese
mountaincat (F. bieti) ............................................. .....96

Fig. 11. Distribution of the Florida
panther (P.c. coryi) ...................................................... 136

Fig. 3. Distribution of the manul (0. ma&) ...............98

Fig. 12. Distribution of the ocelot (L. pardalis) ........ 139

Fig. 4. Distribution of the Asiatic wildcat(F. silvestris,ornata group) ........................................ 100

Fig. 13. Distribution of the bobcat (L. rufus)............. 142

Fig. 5. Distribution of the Eurasianlynx (L. lynx) ..... 102
Fig. 6. Recentdistribution of L. Lynxin Europe......... 104

Fig. 14. Distribution of the pampas
cat (0. colocolo) ..............~........................................... 145

Fig. 7. Distribution of the Iberian lynx
(L. pardinus) ................*................................ ............... 107

Fig. 15. Distribution of three speciesof pampas
cat accordingto Garcia-Perea..................................... 145

Fig. 8. Population structure of the Iberian
lynx in Spain ...............................................*............... 108

Fig. 16. Distribution of the jaguarundi(H. yaguarondi) ........................................................... 147

Fig. 9. Distribution of the Europeanwildca.t(F. silvestris, silvestris group)..................................... 111

Part II. Major Issuesin Cat Conservation
Chapter 1. Cats and Habitat Loss

Table 1. Distribution andpopulation estimatesfor............................92
snow leopard.............

Fig. 1. Humandisturbanceto natural vegetation...... .151

Table 2. Protected areasfor snow leopardsbY
range stateand size............ ..*........................ .................94

Fig. 2a. Global habitat map: closedforest andwoodland .....................................................................156

Table 3. Comparative importance(%) of different
causesof mortality amonglynxes in Spain ............... .109

Fig. 2b. Global habitat map: open or interrupted
woodland..................................................................... 157
Fig. 2c. Global habitat map: settledareas.................158
Fig. 2d. Global habitat map: grassand shrub
complexes.......*............................................................ 159

333

Appendix

7. List of Maps,

Figures,andTablesPage

Page

Fig. 2e. Global habitat map: tundra and desert......... 160

Chapter 4. Trade

Fig. 2f. Global habitat map: major wetlands............ 161

Fig. 1. Cat fur fashionsin the 1960sdisplayedin
Animals magazine,London.................................... .....224

Table 1. Key to global habitat types .......................... 152

Fig. 2. Trade in skinsof the Felidae, 1978-1988.......225

Table 2. Cat speciesoccurrencein major
world habitat types ......................................................154

Fig. 3. Predicted responseto harvestof leopard
populationsin southernand northern Africa ..............228

Table 3. Habitat specialists:catsassociatedwith six or fewer habitat types .................................... 163

Fig. 4. Annual harvest and averagepelt pricefor the lynx in Canada.................................................234

Table 4. Key habitat types for vulnerable
cat species.................................................................... 163

Fig. 5. Annual harvest and averagepelt pricefor the bobcat in the U.S. ............................................234

Table 5. Estimatesof tropical forest area
anddeforestationby ecological zone.......................... 164

Table 1. CITES-reported international trade incat skins, 1976-1990....................................................226

Table 6. Estimatesof tropical forest area
anddeforestationby geographicsub-region...............165

Chapter 5. Cats in Captivity

Table 7. Vulnerable cat speciesstrongly or
significantly associatedwith tropical moist
anddry forest............................................................... 166

Fig. 1. Evolution of zoos............................................245
Fig. 2. Representationof felid speciesin zoo
collectionsin December, 1992....................................247

Table 8. Percentagechangein global areaof
major world habitat types since 1980........................ .168
Table 9. Wildernessareaby geopolitical region ....... 169

Table 1. World zoo associations................................246

Table 10. Protected areaby geopolitical region ........ 173

Table 2. Cat speciesnot representedin ISIS zoos .....248

Table 11. Protected areafor cat species(in order of vulnerability) ............................................ 178

Table 3. CITES-reported international trade inlive cats 1976-1990...................................................*249

Chapter 3. Research

Table 4. Animals of unknown subspecies/Origin
in ISIS zoos in 1989....................... ..*.......................... 250

Fig. 1. Pumatrack identification quiz ........................205

Table 5. Captive propagationstatusof cat species....251

Fig. 2. Track measurements
for individual
identification of pumas...........*............*....................... 205

Table 6. Structure of an AZA SpeciesSurvival
Plan (SSP) Master Plan...............................................252

Table 1. Researcheffort for cat species..................... 197

Table 7. Captive breedingprogramsand
studbooksfor felid species..........................................253
Table 8. A Global Captive Action Plan for Felids.....255

Table 2. Speciesfor which fewer than threeadequatestudiesof natural history have beendone ... 198

Table 9. North American 1993-1994Regional
Collection Plan for Felids............................................257

Table 3. Genetic research:questions,applicabletechniques,and examplesof studiesof felids.............212

Table 10. Australasian1993-1994Regional
Collection Plan for Felids............................................258

Table 4. Ratiosof effective population size toactual population size (Ne/N) calculatedfor
cat populations.............................................................214

Table 11. Goalsof the Tiger Global
Animal Survival Plan ..................................................258

Table 5. Exposureto infectious micro-pathogens
documentedin wild cat populations...........................217
Appendix 2.............................315
Fig. 1. Postmortemmeasurements

334

References

References
Alcala, A.C. and Brown, W.C. 1969. Notes on the food
habits of three Philippine wild mammals. Silliman J.
16: 91-94.
Aldama, J.J., Beltran, J.F. and M. Delibes. 199 1. Energy
expenditure and prey requirements of free-ranging
Iberian lynx in southwestern Spain. J. Wildl. Manage.
55(4): 635-641.
Alekperov,
H.M.,
Yerofeyeva,
S.N. and I.K.
Rakhmatulina. 1977. [Present status of some mammalian species in Azerbaijan.] Pp. 28-34 in [Rare mammals of the U.S.S.R. fauna.] Nauka, Moscow (inRussian).
Alho, C.J., Lather, T.E. and H.C. Goncalves. 1988.
Environmental degradationin the Pantanalecosystem.
Bioscience38: 164-171.
Aliev, F. 1973. The Caucasianblack cat, Felis silvestriscaucasicaSatunin, 1905.Saugetierk.Mitt. 22: 142-145.
Allayarov, A.M. 1963. [Information on the ecology andgeographical distribution of the spotted cat in
Uzbekistan.] Voprosy Biologii i Kraevoi Meditsiny 4:
3 15-321(in Russian).
-. 1964. [Information on the ecology and geographicaldistribution of the jungle cat in Uzbekistan.] [Uzbek J.
Biol.] 8(2): 46-50 (in Russian).
Allen, A. 1987. The relationship between habitat andfurbearers. Pp. 164- 179 in M. Nowak, J. Baker, M.
Obbard and B. Malloch, eds. Wild furbearer management and conservation in North America. Ministry of
Natural Resources,Toronto, Ontario.
Allen, G.M. 1938. The mammalsof China and Mongolia.
Natural history of Central Asia, Vol. XI, Pt. 1.
American Mus. Nat. Hist., New York.
- . 1939. A checklist of African mammals.Bull. Mus.
Camp.2’001.Ham).83: l-763.
Allen, G.O. 1919. Caracal (Felis caracal) and hunting
leopard (Cynailurus jubatus) in Mirzapur, Uttar
Pradesh.J. Bomb. Nat. Hist. Sot. 26.
Allen, J.A. 1903.A new deeranda new lynx from the stateof Sinaloa,Mexico. Bull. Am. Mus. Nat. Hist. 19: 6 14.
- . 1919.Severtzow’s classification of the Felidae. Bull.
Am. Mus. Nat. Hist. 41: 335-340.
Allendorf, F.W. 1983.Isolation geneflow and geneticdifferentiation among populations. Pp. 5 l-65 in C.M.
Schonewald-Cox, S.M. Chambers,B. MacBryde and
W.L. Thomas, eds. Geneticsand conservation: a ref-Abbadi, M. 1992. Israel’s elusive feline: sand cats. Cat
News 18: 15- 16. Bougy-Villars, Switzerland.
Abramov, V.K. and Pikunov, D.G. 1974. [Leopards in thefar east of the U.S.S.R. and their protection.] [Bull.
Moscow Sot. for Nature-Amateur.
Sec. Biol.] 79(2):5-15 (in Russian).
Abu-1 Fazl. 17th century. The Akbar Nama, Vol III.
Translated in New Delhi, 1977.
Achuff, P.L. and Petocz, R. 1988. Preliminary resource
inventory of the Arjin Mountains Nature Reserve,
Xinjiang, People’s Republic of China. Unpubl. project
report, WWF, Gland, Switzerland.
Ackerman, B.B. 1982. Cougar predation and ecologicalenergetics in southern Utah. M.S. thesis, Utah State
Univ., Logan.
-, Lindzey, F.G. and T.P. Hemker. 1986. Predictive energetics model for cougars. Pp. 333-352 in S.D. Miller
and D.D. Everett, eds. Cats ofthe world: biology, conservationand management. National Wildlife
Federation, Washington, D.C.
Adamson, G. 1986. My pride and joy. Collins Harvill,
London.
Adamson, J. 1960. Born free. Collins Harvill, London.
- . 1969. The spotted sphinx. Harcourt, New York.
Ahlborn, G. and Jackson, R.M. 1988. Marking in freeranging snow leopards in west Nepal: a preliminary
assessment. Pp. 24-49 in H. Freeman, ed. Proc. fifthinternational snow leopard symposium. Wildlife
Institute of India and International Snow Leopard Trust,Seattle.
Ahmad, A. 1994. Protection of snow leopards through grazier communities-someexamples from WWFPakistan’s projects in northern areas. In J.L. Fox andDu Jizeng, eds. Proc. seventh international snow leopard symposium. International Snow Leopard Trust,Seattle.
Airumyan, K.A. and Gasparyan, KM. 1977. [Rare ungulates and carnivores in Armenia.] Pp. 35-42 in [Raremammals of the U.S.S.R. fauna.] Nauka, Moscow (inRussian).
Akin, A. 1989. Anatolian leopard on the brink. Cat News
11: 10. Bougy-Villars, Switzerland.
- . 1991. The status of the leopard in Turkey. Pp. 7-10 in
A. Shoemaker, ed. 1990 Internatl. leopard studbook.
Riverbanks Zoological Park, Columbia, South Carolina.

335

References

erence for managing wild animal and plan populations.
Benjamin Cummings, London.
- and Leary, R.F. 1986. Heterozygosity and fitness in natural populations of animals. Pp. 57-76 in M. Soul& ed.
Conservation biology: the science of scarcity anddiversity. Sinauer, Sunderland, Massachusetts.
Alvarez de1 Toro, M. 1952. [Forest animals of Chiapas.]
Ediciones de1 Gobierno de1 Estado, Tuxtla Gutierrez,
Chiapas (in Spanish).
Alvarez, Ken. 1993. Twilight of the panther: biology,
bureaucracy, and failure in an endangered species program. Myakka River Publishing, Sarasota FL., USA.
Alvi, M.A. and Rahman, A. 1968. Jahangir the naturalist. National Institute of Sciences of India, New Delhi.
Ammann, K. 1993. Close encounters of the furred kind.
BBC Wild&e 1 l(7): 14-15.
Ancient Forest International. 1990. Into the Chilean forest.
News of Old Growth 1:1.
Anderson, A. 1983. A critical review of literature on puma
(Felis concolor). Special report 45, Colorado State
Division of Wildlife.
Anderson, D. 1977. Gestation period of Geoffroy’s cat(Leopardus geoflroyi) bred at Memphis Zoo. Int. Zoo
Yearb. 17: 164-l 66.
Anderson, E.M. 1987. A critical review and annotated bibliography of literature on the bobcat. Special report
62, Colorado StateDivision of Wildlife.
Anderson, J.L. 1981. The re-establishmentand management of a lion Panthera leo population in Zululand,South Africa. Biol. Conserv. 19: 107-117.
- . 1992. Re-establishinglarge predators.Reintroduction
News4: 9.
Annenkov, B.P. 1990.The snowleopard (Uncia uncia) in
the Dzungarsky Alatau. Intl. Ped. Book of Snow
Leopards6: 21-24.
Anonymous. 1971. Leopard skin coatslook better on leopards.Animals Magazine (later BBC Wildltye) 352-363.
- . 1974. Symposiumon the contribution of zoos to theconservationof nature held at the Zoological Gardens,
Regent’sPark, London, 8 May 1974. Unpubl. report,
World Wildlife Fund, Morges, Switzerland.
- . 1976. [Ingredients of ChineseMedicine 2.1 ResearchandDevelopmentCenter for ChineseHerbal Medicine,
Taipei (in Chinese).
- . 1978.The internationaltrade in Felidae 1977.Pp. 264293 in 1978Annual report of the CITES Secretariat.
Lausanne,Switzerland.
- . 1979.Frankfurt fur fair. TRAFFIC Bulletin 1(3/4): 8.
- 1984.Cat skinsseizedin Thailand. TRAFFZCBulletin‘6(l): 14.
- . 1985. [Red Data Book of the Turkmen Republic.]
Ashkabad (in Russian).
- . 1986. Cat skins on sale in Lhasa. Cat News 4: 11.
Bougy-Villars, Switzerland.

- . 1987a. Cats in Xinjiang nature reserve, China. Cat
News6: 33. Bougy-Villars, Switzerland.
- . 1987b.Man-eating lions in Tanzania. Cat News7: 22.
Bougy-Villars, Switzerland.
- . 1987~.Exports of medicine hit record. China Daily(Beijing), 15 December.
- . 19876. EEC import restrictions. TRAFFIC Bulletin
8(4): 57-58.
- . 1987e. Silk road leads to snow leopard pelts. SnowLine 11: 7.
- . 1988. Hong Kong gourmets frustrated. Wildl$e
Bulletin (WWF Hong Kong) 6: 2.
- . 1989a.U.S. identifies stateseligible to export bobcat.
TRAFFIC USA Newsletter9(2): 7.
- . 1989b.The easternpuma: evidencecontinuesto build.
Newsletterof the Intl. Sot. Crypto.zoology g(3): l-8.
- . 1989~.Leopardsin Turkey. Cat Newsi0: 8. BougyVillars, Switzerland.
- . 1989d. World record jaguars claimed in Bolivian
Pantanal.Cat News 10: 9. Bougy-Villars, Switzerland.
-. 1990a.Relieving poverty in a resourcesscarce economy. The World Bank Report on Nepal.
- . 1990b.Chinesemedicine threatensAsia’s last tigers.
Cat News 13: 7. Bougy-Villars, Switzerland.
- . 1990~. Rusty-spotted cat photographed in Gir Lion
Sanctuary. Cat News 13: 15. Bougy-Villars,
Switzerland.
- . 1990d.French hunterscall for lynx control. Cat News
13: 2 1. Bougy-Villars, Switzerland.
- . 1990e. Cat news from Djibouti. Cat News 13: 21.
Bougy-Villars, Switzerland.
- . 1990f. Eco-development for India’s tiger areas.Cat
News 14: 10. Bougy-Villars, Switzerland.
- . 1990g.Resultsof a tiger censusin India. Cat News 13:
3. Bougy-Villars, Switzerland.
- . 1991a. Concern over proposed lion release in
Botswana. Cat News 14: 12. Bougy-Villars,
Switzerland.
- . 199lb. Neutering urged for controversial lions in
Botswana. Cat News 15: 6-7. Bougy-Villars,
Switzerland.
- . 199lc. Four ocelotsin a day. Cat News 15: 16. BougyVillars, Switzerland.
- . 1991d.The subspeciesquestion. Cat News 15: 17-18.
Bougy-Villars, Switzerland.
- . 199le. Lynx managementrecommendationsfor priority species.In Management recommendationsfor
Washington’s priority
habitats and species.
WashingtonDept. of Wildlife, Olympia.
- . 1991f. Tigers sighted in northeast China. Cat News
14: 9. Bougy-Villars, Switzerland.
- . 1992a.Iriomote Island: test casefor Japanesepolicy
on biological diversity. Japan Environment Monitor
April 30: 6-9.

336

References- . 1994~. Range countries set up Global Tiger Forum.
Cat News 20: 2-3. Bougy-Villars, Switzerland.
- . 19944. Breeding tigers for profit: ‘wise-use’ or misuse? Bangkok Poyt 23 Nov., Bangkok, Thailand.
- . 1994e. Disease strikes Serenget lions. Cat News 21
Bougy-Villars, Switzerland.
- . 1994f. Records of tiger hunting. Cat News 2 1, Bougy
Villars, Switzerland.
Ansell, W.F.H. and Dowsett, R.J. 1988. Mammals o.
Malawi. Teendine Press, Cornwall.
Anstey, S. 1991. Large mammal distribution in Liberia:
the findings of a preliminary national survey 19891990. Unpubl. report, WWF, Gland, Switzerland.
- . 1992. Angola: environment status quo assessment
report. Unpubl. report, IUCN, Harare.
Aranda, J.M. 1990. [The jaguar (Panthera onca) in the
Calakmul Reserve: morphometrics, food habits, andpopulation density.] M.S. thesis, Universidad National,
Herdia (in Spanish).
- . 199 1. Wild mammal skin trade in Chiapas, Mexico.
Pp. 174-177 in J.G. Robinson and K.H. Redford, eds.
Neotropical wildlife use and conservation. Univ. of
Chicago Press, Chicago.
Arita, H.T., Robinson, J.G. and K.H. Redford. 1990.
Rarity in neotropical forest mammals and its ecological correlates. Conserv. Biol. 4(2): 18 l- 192.
Arra, M.A. 1974. [Distributionof Leo onca (L) in
Argentina (Carnivora, Felidae).] Neotropica 20(63):156- 158 (in Spanish).
Arrighi, J. and Salotti, M. 1988. [The wildcat (Felis silvestris Schreber, 1777) in Corsica: confirmation of its
presence and taxonomic approach.] Mammalia 52( I):
123-125 (in French).
Artois, M. 1985. [Spatial and temporal utilization patternsof the red fox (Vulpes vulpes) and forest wildcat (Felis
silvestris) in Lorraine.] Gibier Faune Sauvage 3: 3357 (in French).
- and Remond, M. 1994. Viral diseases as a threat to freeliving wildcats (Felis silvestris) in Continental Europe.
Vet. Record 134.
Ashman, D., Christensen, G.C., Hess, M.C., Tsukamoto,
G.K. and M.S. Wichersham. 1983. The mountain lion
in Nevada. Unpubl. report, Nevada Dept. Wildl., Reno.
Avenant, N.L. 1993. The caracal, Felis caracal Schreber
1776 as a predator in the West Coast Strandveld. M.S.
thesis, Univ. Stellenbosch, Stellenbosch.
Avise, J.C. and Ball, R.M., Jr. 1990. Principles ofgenealogical concordance in species concepts and biological taxonomy. Oxford Surveys in Evolution Biol.
7: 45-67.
Aymerich, M. 1982. [Comparative study of the diet of thepardel lynx (Lynx pardina Temminck, 1824) and thewildcat (Felis silvestris Schreber, 1777) in the central
Iberian peninsula.] Mammalia 46: 5 15-521 (in French).

- . 1992b. The situation, conservation needs and reintroduction of lynx in Europe. Proc. symp. 17- 19 October,
Neuchatel. Council of Europe, Strasbourg.
- . 1992~. [Felids of Venezuela.] [Foundation for the
Study of Physical, Mathematical and Natural Sciences(FUDECI)], Caracas (in Spanish).
- . 1992d. 1992 annual report. Nyae Nyae Development
Foundation of Namibia, Windhoek.
- . 1992e. Extensive
wildlife
trade in Vietnam.
International Primate Protection League Newsletter
19(l): 3-5.
- . 1992f. Leopard problems. Cat News 16: 8. BougyVillars, Switzerland.
- . 1992g. Rampant skin trade in India. Cat News 16: 25.
Bougy-Villars, Switzerland.
- . 1992h. Collapse in fur sales reported. Cat News 16: 27.
Bougy-Villars, Switzerland.
- . 1992i. Poaching for bones threatens world’s last tigers.
Cat News 17: 2-3. Bougy-Villars, Switzerland.
- . 1992j. Clouded leopard reports. Cat News 17: 11.
Bougy-Villars, Switzerland.
- . 1992k. The Overseas Chinese: a driving force. The
Economist 18 July: 21-24.
- . 19921. Saving the tiger: in the farmyards of the night.
The Economist 14 Mar: 10 1.
- . 1993a. The Amur tiger: problems concerning preservation of the population. Resolutions of the international symposium on the Amur tiger, 11 March 1993,
Wildlife Institute, Khabarovsk, Russia.
- . 1993b. Russia and China set quotas for lynx exports.
Cat News 19: 25. Bougy-Villars, Switzerland.
- . 1993~. The status and conservation of Venezuela’scats. Cat News 18: 18- 19. Bougy-Villars, Switzerland.
- . 1993d. Vegetation classification: report of the UNEPHEM/wCMC/GCTE
preparatory
meeting,Charlotteville, Virginia, 24-26 Jan 1993. Earthwatch
Global Environment Monitoring Systems (GEMS)
report no. 19, UNEP.
- . 1993e. Introduced lioness and cubs shot in Tulireserve, Botswana. Cat News 18: 12. Bougy-Villars,
Switzerland.
- . 1993f. Arabian leopards shot in United Arab Emirates.
Cat News 19: 19-20. Bougy-Villars, Switzerland.
- . 1993g. Russian poachers hit Lazovskiy reserve. Cat
News 18: 5-6. Bougy-Villars, Switzerland.
- . 1993h. Efforts grow to save the Siberian tiger. Cat
News 19: 4-5. Bougy-Villars, Switzerland.
- . 19931. End of the road: let’s accept that there is noway back for Asia’s big cats. Asiaweek 5 May: 35-39.
- . 1994a. Europe’s introduced lynx in peril? Cat News
20: 19. Bougy-Villars, Switzerland.
- . 1994b. Canada lynx added to Washington state
Threatened List. Cat News 20: 21. Bougy-Villars,
Switzerland.

337

References

Aziz, T. 1994. Melghat Tiger Reserve on the chopping
block. WWF India Quarterly Jan-Mar.
Baikov, N. 1936. Big game hunting in Manchuria. Engl.
translation, London.
Bailey, T.N. 1972. Ecology of bobcats with special reference to social organization. Ph.D. thesis, Univ. of
Idaho, Moscow.
- . 1974. Social organization in a bobcat population. J.
WiZdl. Manage. 38: 435446.
- . 1979. Den ecology, population parameters and diet ofeastern Idaho bobcats. Bobcat Res. Conf, Natl. Wildl.
Fed. Sci. Tech. Ser. 6: 62-69.
- . 198 1. Factors of bobcat social organization and somemanagement implications. Pp. 984-1000 in J.A.
Chapman and D. Pursley, eds. Proc. Worldwide
Fur-bearer Conf Frostburg, Maryland.
- . 1993. The African leopard: a study of the ecology andbehavior ofa sohtaryfelid. ColumbiaUniv. Press,NewYork.
- 7 Bangs, E.E., Portner, M.R., Malloy, J.C., and R.J.
McAvinchey. 1986. An apparent overexploited lynx
population on the Kenai Peninsula, Alaska. J. Wildl.
Manage. 50: 279-290.
Baker, J.A. and Dwyer, P.M. 1987. Techniquesfor commercially harvesting furbearers. Pp. 970-995 in M.
Novak, J. Baker, M. Obbard, and B. Malloch, eds.Wild
furbearer management and conservation in North
America. Ministry of Natural Resources, Toronto,
Ontario.
Baker, L.A. 1991. Feeding ecology of reintroduced bobcatson CumberlandIsland, Georgia. M.S. thesis,Univ.
Athens, Georgia.
Balharry, D. and Daniels,M. 1993.Wildcat recovery programmein Scotland: wildcat survey 1983-1987. Pp.
IO2- 110 in Proc. seminar on the biology and conservation of the wildcat (Felis silvestris), Nancy, France,
23-25 September1992.Council of Europe, Strasbourg.
Ballou, J.D. 1992.Potential contribution of cryopreserved
germplasmto the preservationof geneticdiversity andconservation of endangered species in captivity.
Cryobiology 29: 19-25.
Banerjee, L.K. 1984. Vegetation of somecat habitats in
India. Unpubl. report, Botanical Survey of India, Dept.
of the Environment, Howrah, Calcutta.
Banfield, A.W. 1974. The mammalsof Canada. Univ.
Toronto Press,Toronto. ruf Banks, E. 1949.Bornean
mammals.Kuching Press,Kuching.
Bannikov, A.G. 1954. [Mammals of the Mongolian
Peoples’ Republic.] Nauka, Moscow (in Russian).
- and Sokolov, V.I. 1984. [Red Data Book of the
U.S.S.R.,2d edn.] LesnayaPromyshlennost,Moscow.
Barber, G., Horne, L., Mason, M. and S. Perkins. 1984.
NanseiShotoExpedition, Japan. Unpubl. report, WWF
Japan,Tokyo.

Barnes,L. 1989.The overt illegal@- trade in Kathmandu,
NepaE.Unpubl. report, InvernessResearchAssociates,
Inverness,California.
Barnes, R. 1976. Breeding and hand-rearing of the marbled cat. Int. Zoo Yearb. 16: 205-208.
Barton, N.H. and Hewitt, G.M. 1985. Analysis of hybrid
zones.Ann. Rev. Ecol. Syst. 16: 113.
-. Adaptation speciationand hybrid zones.Nature 341:
497-503.
Basilio, A. 1962.La vida animal en la Guinea espanola.
Description y vida de 10sanimalesen la selva tropicalafricana, 2d edn. [Animal life in Spanish Guinea:
descriptionand life of African tropical forest animals.]Inst. EstudiosAfricanos, Madrid (in Spanish).
Beale, D.M. and Smith, A.D. 1973. Mortality of pronghorn antelopefawnsin westernUtah. J. Wildl. Manage.
37: 343-352.
Beardmore, J.A. 1983. Extinction, survival, and genetic
variation. Pp. 125-151 in C.M. Schonewald-Cox,S.M.
Chambers, B. MacBryde and W.L. Thomas, eds.
Geneticsand conservation: a referencefor managing
wild animal and plant populations. BenjaminCummings,London.
Beasom,S.L. and Moore, R.A. 1977. Bobcat food habit
responseto a changein prey abundance.SouthwestNat.
21: 451-457.
Beebe, W. 1925.Ecology of Kartabo. Zoologica 6(l): l193.
Beier, P. 1991. Cougar attacks on humansin the UnitedStatesandCanada.Wildl. Sot. Bull. 19: 403-412.
- . 1993.Determining minimum habitat areasandhabitat
corridors for cougars.Conserv.Biol. 7(l): 94-108.
- and Barrett, R.H. 1990-1992.Orange county cooperative mountain Zion study. Unpubl. quarterly reports,
Department of Forestry and Resource Management,
Univ. of California, Berkeley.
Belden, R.C. and McCown, J.W. 1993. Florida panthercaptive breeding/reintroduction feasibility study:
annual performance report. U.S. Fish and Wildlife
Service, Gainesville, Florida.
Belousova, A.V. 1993. Small Felidae of EasternEurope,
Central Asia, and the Far East: survey of the state of
populations.Lutreola 2: 16-21.
Belovsky, G.E. 1987.Extinction modelsand mammalianpersistence.Pp. 35-58 in M.E. Soul& ed. Viable populations for conservation. Cambridge Univ. Press,
Cambridge.
Beltran, J.F. 1988.[Ecology and spatio-temporalbehaviorof the Iberian lynx (Lynx pardina T.) in the Doiiana
National Park.] Ph.D. thesis,Univ. de Sevilla, Seville(in Spanish).
- 7Aldama, J. and Delibes, M. 1987. Ecology of the
Iberian lynx in Dofiana, SW Spain. Trans. Int. Union
GameBiologists 18.

338

References-

and Delibes, M. 199 1. Feeding ecology of the Iberian
lynx in Dofiana during a drought period. Donana Acta
Vertebrata 18(l): 113-122.
- and Delibes, M. 1993. Physical characteristics
ofIberian lynxes (Lynx pardinus) from Donana, southwestern Spain. J. Mamm. 74(4): 852-862.
-, San Jose, C., Delibes, M. and F. Braza. 1985. An
analysis of the Iberian lynx predation upon fallow deerin the Coto Dofiana, SW Spain. Trans. Int. Union Game
Biologists 17: 961-967.
Berg, W.E. 1979. Ecology of bobcats in northern
Minnesota. Bobcat Res. Conf, Natl. Wildl. Fed. Sci.
Tech. Ser. 6: 55-61.
Berger, J. and Wehausen, J.D. 1991. Consequences of a
mammalian predator-prey disequilibrium in the GreatBasin Desert. Conserv. Biol. 5(2): 244-248.
Bergerud, A. 1983. Prey switching in a simple ecosystem.
Sci. Am. 249(6): 130- 141.
Bernard, R.T.F. and C.T. Stuart. 1987. Reproduction of the
caracal Felis caracal from the Cape Province of South
Africa. S. Afr. J. 2001. 22(3): 177- 182.
Bernhart, F. 1990. [Studies of the activity pattern of thelynx in Switzerland.] Dipl. thesis, Univ. Bern, Bern (inGerman).
Bernier, FranCois. 1670. Histoire de la derniere revolution des &tats du Grand Mogul. [Travels to the Mogul
Empire, 1656- 1668.1 Rev. Engl. transl., Archibald
Constable, London, 189 1.
Berrie, P. 1973. Ecology and status of the lynx in interior
Alaska. Pp. 4-41 in R.L. Eaton, ed. The world’s catsvol. 1: ecology and conservation. World Wildlife
Safari, Winston, Oregon.
Berry, H.H. 1981. Abnormal levels of disease and predation as limiting factors for wildebeest in the Etosha
National Park. Madoqua 11: 242-253.
- . 1991a. Namibia’s seal-eating lions in dangers. Cat
News 14: 10, Bougy-Villars, Switzerland.
- . 1991b. Last Skeleton Coast lions killed by farmers. Cat
News 15: 7-8, Bougy-Villars, Switzerland.
- . 199 lc. Large scale commercial wildlife utilization:
hunting, tourism and animal production in Namibia. Pp.
45-49 in J.G. Grootenhuis, S.G. Njuguna and P.W. Kat,
eds. Wildlife research for sustainable development:
Proc. con& 22-26 April 1990. Kenya Agricultural
Research Institute, Kenya Wildlife Service, andNational Museums of Kenya, Nairobi.
Bertram, B.C.R. 1975a. The social system of lions. Sci.
Am. 232: 54-65.
- . 1975b. Social factors influencing reproduction in wildlions. J. Zool., Lond. 177: 463-482.
- . 1976. Kin selection in lions and in evolution. Pp. 28 l301 in P.P.G. Bateson and R.A. Hinde, eds. Growing
points in ethology. Cambridge Univ. Press, Cambridge.
- . 1978. Pride of lions. Dent, London.

-. 1979. Studying predators, 2d edn. African

Wildlife
LeadershipFoundation, Nairobi.
- . 1982. Leopard ecology as studied by radio tracking.
Symp.2001.Sot. Lond. 49: 341-352.
Berwick, S. 1974. The community of wild ruminants in
the Gir forest ecosystem, India. Ph.D. thesis, Yale
Univ., New London.
- . 1976. The Gir Forest: an endangered ecosystem.
American Scientist64: 28-40.
Bewick, T. 1807. A general history of the quadrupeds.
Bewick andHodgson,Newcastle-upon-Tyne.
Bhattacharyya, T. 1988.A report on thepilot survev on thestatus and distribution offishing cat (Felis viverrina)
in Howrah District of West Bengal. Unpubl. report,
Indian Society for Wildlife Research,Calcutta.
- . 1992.A brief note on someobservationson the breeding biology of the fishing cat (Felis viverrina).
Tigerpaper 19: 20-2I.
Bigourdan, J. and Prunier, R. 1937. [Wild mammalsofWest Africa and their surroundings.] . Montrouge,
France (in French).
Biquand, S. 1990.Short review of the statusof the Arabianleopard, Panthera pardus nimr, in the Arabian peninsula. Pp. 8- 10 in A. Shoemaker, ed. 1989 Znternatl.
leopard studbook. Riverbanks Zoological Park,
Columbia, South Carolina.
Birkeland, K.H. and Myrberget, S. 1980. The diet of thelynx Lynx lynx
I in Norway. Fauna Now. I: 24-28.
Bisbal, F.J. 1986. Food habits of someneotropical carnivores in Venezuela (Mammalia, Camivora). Mammalia
50(3): 329-339.
- . 1989. Distribution and habitat associationof the carnivores in Venezuela.Pp. 339-362in K.H. Redford andJ.F. Eisenberg,eds.Advances in neotropical mammalogy. Sandhill CranePress,Gainesville, Florida.
Biswas, B. and Ghose, R.K. 1982. Progress report I on
pilot survey of the WWF-India/Zoological survey ofIndia collaborative project on the statussurvey of thelessercats in easternIndia. Unpubl. report, Zoological
Survey of India, Calcutta.
- 7andGhosal,D.K. 1985.Progressreport 2 onpilot survey of the WWF-India/Zoological survey of India collaborative project on the statussurvey of the lessercats
in easternIndia. Unpubl. report, Zoological Survey ofIndia, Calcutta.
Bjarvall, A. 1992. Lynx and reindeer management in
Sweden.Pp. 40-42 in The situation, conservationneeds
and reintroduction of lynx in Europe. Proc. Symp. 1719 October, Neuchatel.Council of Europe, Strasbourg.
Blanford, W.T. 1891. Thefauna of British India, including
Ceylon and Burma: Mammalin. London.
Blankenship, T.L. and Swank, W.G. 1979. Population
dynamic aspectsof the bobcat in Texas. Bobcat Res.
Con&,Natl. Wildl. Fed. Sci. Tech Ser. 6: 116-122.

References

Blomqvist, L. 1978. First report on the snow leopard studbook (Panthera uncia) and the 1977 world register.
Intl. Ped. Book of Snow Leopards 1: 22-34.
- 7ed. 1990. International pedigree book ofsnow leopards, VOL.6. Helsinki Zoo, Finland.
- and Sten, I. 1982. Reproductive biology of the snow
leopard, Panthera uncia. Intl. Ped. Book of Snow
Leopards 2: 71-79.
Bluett, R.D. and Tewes, M.E. 1988. Evaluation ofbobcat
harvest relative to estimated population size and habitat base in Texas, 1978-1986. Caesar Kleberg Wildlife
Research Institute, Univ. of Texas, Kingsville.
Blum, L.G. and Escherisch, P.C., eds. 1979. Bobcat
research conference proceedings. Nat. Wildl. Fed.,
Tech. Rep. Ser. 6.
Boesch, C. 199 1. The effects of leopard predation ongrouping patterns in forest chimpanzees. Behaviour
117(3/4): 220-242.
Boggess, E., Berg, B. and D. Kuehn. 1989. Minnesota registered&r-bearer harvest statistics and population status. Wildlife Section, Minnesota Dept. of Natural
Resources, St. Paul.
Bold, A. and Dorzhzunduy, C. 1976. Reports about theirbis of the southern mountains of the Gobi Altay.
Trudy instituta obshchey i eksperimental ‘nay biologii
11: 27-43 (Ulaan Baator).
Bolgiano, C. 199 1. Of panthers and prejudice. Buzzworm
May/June: 47-5 1.
Bondar, B.N. 1987. [The distribution of the wildcat
throughout the natural zones of the Carpathians.] [Proc.
Congr. AZ&Union Therio. Sot.] 3: 28-30. Nauka,
Moscow (in Russian).
Borner, M. 1977. Leopards in western Turkey. Oryx 14( 1):26-30.
Bothma, J. du P. and Le Riche, E.A.N. 1984. Aspects ofthe ecology and the behaviour of the leopard Panthera
pardus in the Kalahari Desert. Koedoe Suppl. : 259-279.
- . 1986. Prey preference and hunting efficiency of the
Kalahari Desert leopard. Pp. 389-414 in SD. Miller and
D.D. Everett, eds. Cats of the world: biology, conservation and management. National Wildlife Federation,
Washington, D.C.
Bottriell, L.G. 1987. King cheetah: the story of the quest.
EJ Brill, Leiden.
Bourdelle, E. and Babault, G. 193 1. [Note on an unusual
form of felid from the Kivu region.] Bull. Mus. Hist.
nat., Paris 2(3): 294-297 (in French).
Bourliere, E. 1955. Mammals of the world: their life andhabits. Alfred A. Knopf, New York.
Boutin, S., Krebs, C.J., Sinclair, A.R.E. andJ.N.M. Smith.
1986. Proximate causesof lossesin a snowshoehare
population. Can. J. 2001.64: 606-610.
Bowland, A.E. 1993. The 1990/1991 cheetah photographic survey. Unpubl. report, Departmentof Nature

Conservation, Skukuza, Kruger National Park, South
Africa.
-, Mills, M.G.L. and D. Lawson. 1993. Predators andfarmers. EndangeredWildlife Trust, Parkview, South
Africa.
Bowland, J.M. 1990.Diet, homerangeand movementpatterns of servals on farmland in Natal. M.Sc. thesis,
Univ. of Natal, Pietermaritzburg.
Bowns, J.E. 1985.Predation-depredation.Pp. 204-215 in
J. Robersonand F.G. Lindzey, eds.Proc. secondmountain lion workshop.Salt Lake City, Utah.
Boydell, C. 1980.Discrepanciesin CITES trade statistics
between Germany and Brazil, Bolivia, Paraguay and
Uruguay. TRAFFIC Bulletin 2(5/6): 52-53.
Boyle, K.A. and Fendley, T.T. 1987. Habitat suitability
index models:Bobcat. U.S. Fish and Wildl. Ser. Biol.
Rep. 82(10.147).
Braden, K.E. 1982. The geographic distribution of snow
leopards in the U.S.S.R. Intl. Ped. Book of Snow
Leopards3: 25-39.
Bragin, A.P. 1986.Population characteristicsand socialspatialpatterns of the tiger on the easternmacro-slopeof the Sikhote-Alin mountain range, U.S.S.R.M.S. thesis,Pacific Institute of Geography, Vladivostok. Orig.
in Russian,Engl. transl. by author.
- . 1990. A short review of the status of the leopard,
Pantherapardus ciscaucasica(saxicolor), in the southwest U.S.S.R. Pp. 11-12 in A. Shoemaker, ed. 1989
Internatl. leopard studbook. Riverbanks ZoologicalPark, Columbia, South Carolina.
- and Gaponov, V.V. 1989.Problemsof the Amur tiger.
Hunting and Hunting Science,Ott (in Russian).Engl.
transl. by A. Bragin, summaryin Cat News 12: 18-19(1990). Bougy-Villars, Switzerland.
Brand, C.J. and Keith, L.B. 1979.Lynx demographyduring a snowshoe hare decline in Alberta. J. Wildl.
Manage. 43: 827-849.
- 7and C.A. Fischer. 1976. Lynx responsesto changing
snowshoehare densities in central Alberta. J. Wildl.
Manage. 40: 4 16-428.
Brand, D.J. 1989. [The control of caracal (Felis caracal)
and baboons(Papioursinus)in the CapeProvince withthe help of mechanical means.] M.S. thesis, Univ.
Stellenbosch,Stellenbosch(in Afrikaans).
Brander, A. 1923. Wild animals in central India. Natraj,
Dehra Dun ( 1982reprint).
Braiitigam, A. 1989. CITES: a conservation tool, 2d edn.
IUCN/SSC Trade Specialist Group and the Center for
Marine Conservation,Washington, D.C.
Breeden,S. 1989.The happy fisher. BBC Wildlife 7: 238241.
Breitenmoser, U. 1983. Zur Wiedereinburgerung desLuchses(Lynx lynx L.) in der Schweiz. [Reintroduction
and spreadof the lynx in Switzerland.] 2. Forstwes.

340

References

134(3): 207-222.
- . 1991. Eurasian lynx (Lynx lynx). Unpubl. report,
IUCN/SSC
Cat Specialist Group, Bougy-Villars,
Switzerland.
Breitenmoser, U. and Baettig, M. 1992. Wiederansiedlung
und Ausbreitung des Luchses (Lynx lynx) im Schweizer
Jura. [Reintroduction and spread of lynx in the Swiss
Jura Mountains.] Rev. suisse 2001. 99( 1): 163-176 (inGerman).
Breitenmoser, U. and Breitenmoser-Wursten,
C. 1990.
Status, conservation needs and r-e-introduction of thelynx Lynx lynx in Europe. Nature and Environment
Series, No. 45. Council of Europe, Strasbourg.
Breitenmoser, U. and Breitenmoser-Wursten,
C. In prep.
The speciation of the Eurasian lynx (Lynx lynx) and theCanada lynx (Lynx canadensis): evidence from theircomparative ecology.
Breitenmoser,
U. and Haller,
H. 1987. Zur
Nahrungsokologie
des Luchses Lynx lynx im den
Schweizerischen Nordalpen. [Feeding ecology of thelynx in northern Swiss Alps.] 2. Saugetierk. 52: 16%
191 (in German).
- . 1993. Patterns of predation by reintroduced European
lynx in the Swiss Alps. J. Wildl. Manage. 57(l): 144154.
U., Kaczensky,
P., Dotterer,
M.,
Breitenmoser,
Breitenmoser-Wursten,
C., Capt., S., Bemhart, F. and
M. Liberek. 1993a. Spatial organization and recruitment of lynx (Lynx lynx) in a reintroduced population in
the Swiss Jura Mountains. J. Zool., Lond. 23 1: 449-464.
Breitenmoser, U., Slough, B.G. and C. BreitenmoserWursten. 1993b. Predators of cyclic prey: is the
Canada lynx victim or profiteer of the snowshoe hare
cycle? Oikos 66: 55 l-554.
Breitenmoser, U., Breitenmoser-Wursten,
C. and S. Capt.
1994. [The Swiss lynx could disappear again.] Panda
Nouvelles (WWF-France) Feb: 3-5. Summarized in Cat
News 20: 19. Bougy-Villars, Switzerland.
Bridgeford, P.A. 1985. Unusual diet of the lion Pantheralea in the Skeleton Coast Park. Madoqua 14: 187- 188.
Broad, S. 1987. The harvest of and trade in Latin
American spotted cats (Felidae) and otters (Lutrinae).
Unpubl. report, World Conservation
Monitoring
Centre, Cambridge.
- . 1988. Little spotted cat, tiger cat, or oncilla. Pp. 124130 in Significant trade in wildhfe: a review of selected
species in CITES Appendix II, Volume 1: Mammals.
World Conservation Monitoring Centre, Cambridge.
Brocke, R.H. and Gustafson, K.A. 1992. Lynx Lynx
canadensis in New York State. Reintroduction News 4.
Brooks, A.C. 1962. Uganda’s small mammals, II: The
small cats. WiZdZzfeand Sport 3(2): 8- 11.
Brooks, D.M. 1990. An update on the status and abundanceof Paraguayan felids, March 1990. Felid 4(2): 2-4.

-

in the Paraguayan Chaco. Cat News 16:
19-23. Bougy-Villars, Switzerland.
Brown, D.E. 1991. Revival for el tigre? Defenders 66( 1):27-35.
Brown, E.W., Miththapala, S. and S.J. O’Brien. 1993.
Prevalence of exposure to Feline Immunodeficiency
Virus in exotic felid species. J. Zoo and Wildl.
Medicine 24(3): 357-364.
Brown, J. 1992. Progress in fecal steroid analysis to noninvasively monitor reproductive status in felids. Pp. 3335 in D.E. Wildt, J.D. Mellen and U.S. Seal, eds. Felid
action plan, 1991 and 1992: AAZPA Felid Taxon
Advisory Group regional collection plan and IUCN
Captive Breeding Specialist Group globalfelid captive
action plan. National Zoological Park, Front Royal,
Virginia.
Browne, M.W. 1992. Folk remedy demand may wipe out
tigers. New York Times 22 Sept.
Bryant, J.P. 1981. Phytochemical deterrence of snowshoe
hare browsing by adventitious shoots of four Alaskan
trees. Science 213: 889-890.
- 7Wieland, G.D., Clausen, T., and P. Kuropat. 1985.
Interaction of snowshoe hare and feltleaf willow in
Alaska. Ecology 66: 1564-1573.
Bulmer, M.G. 1974. A statistical analysis of the lo-year
cycle in Canada. J. Anim. Ecol. 43: 701-7 18.
Burger, M. 1970. [Observations on the Korean leopard,
Panthera pardus orientalis (Schlegel).] 2001. Gart.
38(6): 328-333 (in German).
Burmeister,H. 1879. [PhysicaE descriptionof theArgentine Republic, Vol. 3 No. I.] Buenos Aires (inFrench).
Bumey, D. 1980. The effects of human activities on cheetahs (Acinonyx jubatus Schr.) in the Mara region of
Kenya. M.S. thesis, Univ. of Nairobi, Nairobi.
Burton, R.W. 1950. Rabies in tigers-two
provedinstances. J. Bombay Nat. Hist. 49: 538-541.
Bush, M., Munson, L., Phillips, L., Allen, M., Kramer, L.
and R. Junge. 1992. A guide to medical/nutritional
management of felids. Pp. 149-174 in D.E. Wildt, J.D.
Mellen and U.S. Seal, eds. Felid action plan, 1991 and1992: AAZPA Felid Taxon Advisory Group regional
collection plan and IUCN Captive Breeding Specialist
Group global felid action plan. National ZoologicalPark, Front Royal, Virginia.
Buttner, K. in press. 10 Jahre WiedereinburgerungderEuropaischen Wildkatze Felis silvestris silvestris
Schreber in Bayem (1984- 1993).
Buttner, K. and G. Worel. 1990. Wiedereinburgerung deseuropaischen Wildkatze in Bayern-ein
Projekt desBundes Naturschutz in Bayern. [ReintroductionofEuropean wildcats in Bavaria-Projectof the Bavarian
League for the Protection of Nature.] WaZdhygiene 18:
169-176 (in German).

341. 1992. FelidsReferencesin an asocial species. Univ. Chicago Press, Chicago.
- and Collins, D.A. 1986. Male cheetahs of the Serengeti.
Nat. Geog. Res. 2 75-86.
- 1987a. Male cheetah social organization and territoriality. Ethology 74: 52-64.
- 1987b. Ecological characteristics of territories of malecheetahs (Acinonyx jubatus). J. Zool., Lond. 211: 89105.
Caro, T.M. and Durant, S.M. 1991. Use of quantitative
analyses of pelage characteristics
to reveal family
resemblances in genetically monomorphic cheetahs. J.
Heredity 82: 8-14.
Caro, T.M., FitzGibbon, C.D. and M.E. Holt. 1989.
Physiological costs of behavioural strategies for malecheetahs. Anim. Behav. 38: 309-317.
-, Holt, M.E., FitzGibbon, C.D., Bush, M., Hawkey,
C.M. and R.A. Kock. 1987. Health of adult free-living
cheetahs. J. Zool., Lond. 212: 573-584.
- and Laurenson, M.K. 1990. Serengeti cheetah project.
Pp. 33-39 in Huish, S. and Campbell, K., eds. Biennial
report of scienttjk results for 1988 and 1989. Serengeti
Wildlife Research Centre, Arusha. jubsub-. 1994. Ecological and genetic factors in conservation:a cautionary tale. Science 263: 485-486.
Caroco, T. and Wolf, L.L. 1975. Ecological determinantsof group sizes of foraging lions. Am. Nat. 109: 343-352.
Carpaneto, G.M. and Germi, F.P. 1989. The mammals in
the zoological culture of the Mbuti Pygmies in northeastern Zaire. Hystrix 1: l-83.
Carter, T.D. 1955. Remarkable age attained by a bobcat. J.
Mammal. 36: 290.
Carvalho, C.T. 1958. [On some mammals of southeastern
Para.] Arquivos de Zoologia 12(5): 121- 132 (inPortugese).
Cat Specialist Group. 1984. Saving the wild cats. Manifesto of the IUCN/SSC Cat Specialist Group, BougyVillars, Switzerland.
Caughley, G.F. 1994. Directions in conservation biology.
J. Animal Ecol. 65: 215-244.
Cavallo, J. 1993. A study of leopard behaviour and ecology in the Seronera Valley, Serengeti National Park.
Serengeti Wildlife Research Centre scientific report
1990-1992: 33-43.
Chakrabarty, Kalyan. 1992. Man-eating tigers. Darbari
Prokashan, Calcutta.
Chakraborty, S. 1978. The rusty-spotted cat, Felis rubiginosa I. Geoffroy, in Jammu and Kashmir. J. Bombay
Nat. Hist. Sot. 75: 478-479.
-, Chakraborty, R. and V.C. Agrawal. Melanism in thejungle cat. J. Bombay Nat. Hist. Sot. 85( 1): 184.
Champion, F. 1927. With a camera in Tiger-Land. Chatto
and Windus, London.
Chavan, S. 1993. Life history information on P.Z. persica.
Estimating the carrying capacity of lions in Gir. In

Bygott, J.D., Bertram, B.C.R. and J.P. Hanby. 1979. Malelions in large coalitions gain reproductive advantages.
Nature 282: 839-841.
Cabrera, A. 19 14. [Iberian fauna: mammals. ] Museo
National de Ciencias Naturales, Madrid (in Spanish).
- . 196 1. [Living felids of the Republic of Argentina.]
Rev. Mus. Argent.
Cienc. Nat. “Bernardino
Rivadavia”, 2001. 6(5): 161-247 (in Spanish).
- and Yepes, J. 1960. [South American mammals.]
Historia Natural, Buenos Aires (in Spanish).
Cabrera, A.L. and Willink, A. 1980. [Biogeography of
Latin America, 2d edn.] Organization of American
States, Programa Regional de Desarrollo Cientifico y
Tecnologico, Washington D.C. (in Spanish).
Cai, G.-Q. et al. 1989a. [Economic fauna in Qinghai].
Peoples’ Press of Qinghai, Xining (in Chinese).
-, Liu, Y. and B. O’Gara. 1989b. Observations of large
mammals in the Qaidam Basin and its peripheral mountainous area in the People’s Republic of China. Can. J.
2001. 68: 202 l-2024.
Caldwell, J. 1984. South American cats in trade: theGerman connection. TRAFFIC Bulletin 6(2): 3 l-32.
Captive Breeding Specialist Group (CBSG). 199 1. Genetic
management considerations for threatened species witha detailed analysis of the Florida panther (Felis concolor coryi). Unpubl. report of a genetic augmentation
workshop sponsored by the U.S. Fish and Wildlife
Service and IUCN/SSC Captive Breeding Specialist
Group. Apple Valley, Minnesota.
-. 1993. Global zoo directory. Apple Valley, Minnesota.
Carbyn, L.N. and Patriquin, D. 1983. Observations onhome range sizes, movements, and social organizationof lynx, Lynx canadensis, in Riding Mountain National
Park, Manitoba. Can. Field Nat. 97: 262-267.
Carey, A.B. and McLean, R.G. 1978. Rabies antibody
prevalence and virus tissue tropism in wild carnivores
in Virginia. J. Wildl. Dis. 14: 487-491.
Carey, J.R. and Keith, L.B. 1979. Reproductive change in
the IO-year cycle of snowshoe hares. Can. J. 2001. 57:
375390.
Carman, R.L. 1984. [Southern limit of the distribution of
the jaguar (Leo onca) in the 18th- 19th centuries.] Rev.
Mus. Argent. Cienc. Nat. “Bernardino Rivadavia, ”
2001. 13( l-60): 293-296 (in Spanish).
Caro, T.M. 1982. A record of cheetah scavenging in the
Serengeti. Afr. J. Ecol. 20: 213-214.
-. 1989. Determinants of asociality in felids. Pp. 41-74
in V. Standen and R.A. Foley, eds. Comparativesocioecology: the behavioural ecology of humans andother mammals. Blackwell, Oxford.
- . 1993. Behavioral solutions to breeding cheetahs in
captivity: insights from the wild. Zoo BioEogy 12( 1):19-30.
- . 1994. Cheetahs of the Serengeti plains: group living

342

I

Referencesmodern.British Museum, London.
- . 1992.How the wild beastswere tamed.New Scientist,
15February: 41-43.
Cobb, S. 1981. The leopard-problems of an overabundant, threatened, terrestrial carnivore. Pp. 191- 192 in
P.A. Jewel1and S. Holt, eds.Problemsin managementof locally abundant wild mammals.Academic Press,
New York.
Collier, G.E. and O’Brien, S.J. 1985. A molecular phylogeny of the Felidae: immunological distance.
Evolution 39: 473-487.
Collins, N.M., ed. 1990. The last rainforests. Mitchell
Beazley andIUCN, London.
- 7Sayer, J.A. and Whitmore, T.C., eds. 1991. The conservation atlas of tropical forests: Asia and the Pacific.
IUCN, Gland, Switzerland.
Conde,B. and Schauenberg,P. 1969. [Reproductionof theEuropean forest wildcat (Felis silvestris Schreber) in
captivity.] Rev. Suisse2001.76(7) (in French).
- . 1971. [Weight of the European forest wildcat.] Rev.
Suisse2001.78: 295-315 (in French).
- . 1974. [Reproductionof the forest wildcat (F. silvestris
Schreber) in north-eastern France.] Rev. Suisse2002.
81: 45-52 (in French).
- 7 Nguyen, T.T.C., Vaillant, F. and P. Schauenberg.
1972. [Diet of the forest wildcat (Felis silvestris
Schreber) in France.] Mammalia 36: 112- 119 (inFrench).
Conner, M.C., Labisky, R.F. and D.R. Progulske. 1983.
Scent-stationindices asmeasuresof population abundancefor bobcats,raccoons,gray foxes, andopossums.
Wildl. Sot. Bull. 11: 146-152.
Conway, W. 1968. The consumptionof wildlife by man.
Animal Kingdom, June: 2-7.
- . 1980. An overview of captive propagation. Pp. 199208 in M.E. Soule andB.A. Wilcox, eds.Conservation
biology: an evolutionary-ecological perspective.
Sinauer,Sunderland,Massachusetts.
Cooper, J. 1942. An exploratory study on African lions.
Comp.Psychol. Monograph 17(7): l-48.
Cop, J. 1992. Reintroduction of lynx in Yugoslavia. Pp.
60-63 in The situation, conservation needsand reintroduction of lynx in Europe. Proc. symp. 17-I 9
October, Neuchatel. Council of Europe, Strasbourg.
Corbett, G.B. and Hill, J.E. 1993. The mammalsof the
Indo-Malayan region: a systematic review. Oxford
Univ. Pressfor British Museum (Natural History).
Corbett, J. 1944. Man-eaters of Kumaon. Oxford Univ.
Pressedn. 1946, London.
- . 1948. The man-eating leopard of Rudraprayag.
Oxford Univ. Press,New Delhi.
- . 1953.Jungle lore. Oxford Univ. Press,Oxford.
- . 1957. Man-eaters of India. Oxford Univ. Press,
Oxford.

Asiatic lion PHVA and GASP briefing book, Captive
Breeding Specialist Group, Apple Valley, Minnesota.
Chazee, L. 1990. The mammals of Laos and the hunting
Gland,
practices.
Unpubl. report, WWF/IUCN,
Switzerland.
Chepko-Sade, B.D., Shield, W.M., Berger, J., Halpin,
Z.T., Jones, W.T., Rogers, L.L., Rood, J.P. and A.T.
Smith. 1987. The effects of dispersal and social structure on effective population size. Pp. 287-321 in B.D.
Chepko-Sade and Z.T. Halpin, eds. Mammalian dispersal patterns. Univ. Chicago Press, Chicago.
Child, G.S. 1965. Some notes on the mammals of
Kilimanjaro. Tanganyika Notes and Records 64: 77-89.
Chopra, R. 1988. Fur smuggling in Kashmir. Himal 1( 1):26.
Choudhury, A. 1993. The clouded leopard in Assam. Oryx27( 1): 5 l-53.
Christie, P.M. 1993. Australasian 1993-1994 regional collection plan for felids. In AustralasianSpeciesManagement Program regional census and plan.
ASMP, Mosman, New South Wales.
Chundawat, R.S. 1990a. Lynx survey in Nubra valley,
Ladakh. Wildlife Institute ofIndia Newsletter 5(2): 4244.
Chundawat, R.S. 1990b. Habitat selection by a snow leopard in Hemis National Park, India. Zntl. Ped. Book of
Snow Leopards 6: 85-92.
- and Rawat, R.S. 1994. Food habits of snow leopard in
Ladakh, India. In J.L. Fox and Du Jizeng, eds. Proc.
seventh internationalsnow leopard symposium.
International Snow Leopard Trust, Seattle.
- 7 Rodgers, W. and H. Panwar. 1988. Status report onsnow leopard in India. Pp. 113- 120 in H. Freeman, ed.
Proc. fifth international snow leopard symposium.
International Snow Leopard Trust, Seattle.
Chykovany, T.G., Vronsky, N.V., Gigaury, G.N. and G.K.
Ichuaidze. 1990. [The Akhmetskiy Reserve.] Pp. 200210 in [Reserves of the Caucasus.] Moscow (inRussian).
Cisin, C. 1967. EspeciaZZy ocelots. Harry G. Cisin, NewYork.
Clark, A. 1901. Sport in the low-country of Ceylon. Tisara
Prakasakayo,Dehiwela (1971 reprint).
Clark, C.W. 1987.The lazy, adaptablelions: a Markovian
model of group foraging. Anim. Behav. 35: 361-368.
Clark, J.M. 1976.Variations in coat colour genefrequencies and selectionin the cats of Scotland. Genetica 46:
401-412.
Cloudsley-Thompson, J.L. 1984. Introduction. Pp. I- 15
in J.L. Cloudsley-Thompson, ed. Sahara Desert.
PergamonPress,Oxford.
Clutton-Brock, J. 1981. Domesticatedanimalsfrom earlytimes.British Museum (Natural History), London.
- . 1988. The British Museum book of cats ancient and343

References

Socialist Republic. Vol. 1: Vertebrate animals.]
Academy of Sciencesof the Armenian SSR, Yerevan(in Russian).
Daly, R.H. 1990. Arabian leopard Panthera pardus nimr.
Cat News 12: 4. Bougy-Villars, Switzerland.
Danilov, P.I., Rusakov, O.S. and Tumanov, I.L. 1979.
[Predators in northeastern U.S.S.R..] Leningrad:
Nauka (in Russian).
Darwin, C. 1868. The variation of animals and plants
under domestication,vol. 1. JohnMurray, London.
Dathe, H. 1980. The history of wild animal studbooks.
Inter. Zoo Yearb. 20: 485-486.
Daugherty, C.H., Cree,A., Hay, J.M. andM.B. Thompson.
1990.Neglected taxonomy and continuing extinctionsof tuatara (Sphenodon).Nature 347: 177-179.
Davies, G. and Payne, J. 1982.A fauna1 survey of Sabah.
Unpubl. report, WWF Malaysia for the Game Branch,
Forest Dept., Sabah.
Davies,R.G. 1990.Sighting of a cloudedleopard(Neofelis
nebulosa) in a troop of pigtail macaques (Macaca
nemestrina)in Khao Yai National Park, Thailand. Nat.
Hist. Bull. Siam. Sot. 28: 95-96.
Davis, D.D. 1962.Mammals of the lowland rain-forest of
North Borneo.Bull. SingaporeNat. Hist. Mus. 31: l- 129.
Davis, S.J.M. 1987. Thearchaeologyof animals.Batsford,
London.
de Almeida, A.E. 1984. Somefeeding and other habits ofjaguar in the Pantanal. Unpubl. mss.,SaoPaulo.
de Alwis, W.L.E. 1973. Statusof SoutheastAsia’s small
cats. Pp. 198-208 in R.L. Eaton, ed. The world’s cats,vol. I: ecology and conservation. World Wildlife
Safari, Portland, Oregon.
de Meulenaer, T. and Gray, J.,eds. 1992. The control of
wildhfe trade in Greece. TRAFFIC International,
Cambridge,U.K.
de Pienaar,U. 1969.Predator-prey relationshipsamongst
the larger mammals of the Kruger National Park.
Koedoe 12: 108-176.
de Smet, K.J.M. 1989.[Distribution and habitat choice oflarger mammalsin Algeria, with special reference tonature protection.] Ph.D. thesis, Ghent State Univ.,
Belgium. Original in Dutch: Engl. transl. World
ConservationMonitoring Centre, Cambridge.
Delibes, M. 1979. Le lynx dans la peninsule iberique:repartition et regression.[The lynx in the Iberian peninsula: distribution and decline.] Bull. Mens. 08 Nat.
Chasse,n. sp. Sci. tech., Le lynx: 41-46 (in French).
-. 1980.Feedingecology of the Spanishlynx in the Coto
Dofiana.Acta Theriologica 25: 309-324.
- . 1989.Factorsregulatinga naturalpopulationof lynxes.
Pp. 96-99 in Proc. Con&Reintroductionof Predators inProtected Areas, Torino, Italy.
Deutsch, L.A. 1975. ContribuiCao para o conhecimento
da Panthera onca (Linne)--onGa pintada (Mammalia-

Corbett, L.K. 1979. Feeding ecology and social organization ofwildcats (Felis silvestris) and domestic cats
(Felis catus) in Scotland. Ph.D. thesis, Univ. Aberdeen,
Aberdeen.
Corkill, N.L. 1929. On the occurrence of the cheetah(Acinonyx jubatus) in Iraq. J. Bombay Nat. Hist. Sot.
33: 760-762.
Courtney, N. 1980. The tiger: symbol offreedom. Quartet,
London.
Cox, R. 1988. The conservation status of biologicalresources in the Philippines. Unpubl. report by theWorld Conservation Monitoring Center to the International Institute for Environment and Development,
Cambridge.
Crandall, L. 1964. The management ofwild animals in
captivity. Univ. of Chicago Press, Chicago.
Crawshaw,
P.G. Jr. and Quigley, H.B. 1989. Ocelot
movement and activity patterns in the Pantanal Region,
Brazil. Biotropica 21(4): 377-379.
- . 1991. Jaguar spacing, activity and habitat use in a seasonally flooded environment in Brazil. J. Zool., Lond.
223: 357-370.
- . In prep. Notes on jaguar and puma feeding habits in
the Pantanal ofMato Grosso, Brazil, with implications
for their management and conservation.
Crespo, J.A. 1982. [Ecology of a mammal community in
Iguazti National Park, Misiones.] Rev. Mus. Argent.
Cienc. Nat. “Bernardino Rivadavia, ” Ecol. 3(2): 45
162 (in Spanish).
Crovella, S., Montagnon, D. and E. Natoli. 1993. Highly
repeated DNA sequences in European wildcat anddomestic cat (Carnivora). Pp. 82-85 in Proc. seminaron the biology and conservation of the wildcat (Felis
silvestris), Nancy, France, 23-25 September 1992.
Council of Europe, Strasbourg.
Crowe, D.M. 1975a.Aspects of aging, growth and reproduction of bobcats from Wyoming. J. Mammal. 56:
177-198.
Crowe, D.M. 1975b.A model for exploited bobcat populations in Wyoming. J. Wildl. Manage. 39: 408-415.
Cumberland,R. 1993. Evidence of the easterncougar in
New Brunswick. Unpubl. report, Manitoba Dept. of
Natural Resources,Winnipeg, Canada.
Cunningham, A.B. and Zondi, AS. 1991. Useof animal
partsfor the commercialtrade in traditional medicines.
Institute of Natural Resources, Univ. Natal,
Pietermaritzburg.
Currier, M.J.P. 1983.Felis concolor. Mammalian Species
200: l-7.
-, Sheriff, S.L. and K.R. Russell. 1977. Mountain lion
population and harvest near Canon City, Colorado
1974-l 977. Colorado Division of Wildlife Special
Report 42.
Dal’, SK. 1954. [Animal world of the Armenian Soviet344

References

Carnivora). Cruzamento de exemplares pintadas con
melanicos. [Contributions
to the knowledge of thejaguar: cross-breeding of spotted and melanistic forms.]
Ciencias Biol. Secao 5, Zoologica 5: 369-370 (inPortugese).
Dias, A.S. 1966. [Mammals of Mocambique.] V. Jornados
Medico Veterinarias (Nov.), Lourenco Marques (inPortuguese).
Dickson, H. 1949. The Arab of the Desert. Allen and
Unwin, London, U.K.
Diefenbach, D.R. 1992. The reintroduction of bobcats to
Cumberland Island, Georgia: validation of the scentstation survey technique and analysis ofpopulation viability. Ph.D. thesis, Univ. Georgia, Athens.
Dimitri, M. 1972. [The Andean-Patagonian forest region:
general synopsis.] Coleccion cientwca de1 InstituteNational de Tecnologia Agropecuaria 10 (in Spanish).
Dimitrijevic, S. and V. Habijan. 1977. [Spreading andnumbers of the wild cat FeZis silvestris Schreber, 1777
in Vojvodina.] Arh. Biol. Nauka., Beograd 29: 175
178 (in Russian).
Dinerstein, E. and Mehta, J.N. 1989. The clouded leopard
in Nepal. O~.X 23 (4): 199-201.
Dittrich, L. 1979. [Heredity of melanism in jaguars(Panthera onca).] 2001. Garten 49: 9-23 (in German).
Divyabhanusinh. 1984. The origin, range and status of the
Asiatic (or Indian) cheetah or hunting leopard(Acinonyx jubatus venaticus). Pp. 183- 195 in The plightof the cats: Proc. meeting and workshopof theIUCN/SSC Cat Specialist Group at Kanha National
Park, Madhya Pradesh, India, 9-12 April 1984.
Unpubl. report, IUCN/SSC
Cat Specialist Group,
Bougy, Switzerland.
Dobroruka, L.J. 1964. [Hodgson’s leopard, Panthera pardus pernigra Hodgson 1863.1 Zoo/. Gart. 29(2): 61-67(in German).
- 1969. [Further notes on Panthera pardus pernigra‘(Hodgson, 1863).] 2001. Anzeiger 182( l/2): 52-55 (inGerman).
Donaghue, A.M., Howard, J.G., Byers, A.P., Goodrowe,
K.L., Bush, M., Blumer, E., Lukas, J., Stover, J.,
Snodgrass, K. and D.E. Wildt. 1992. Correlation of
sperm viability with gamete interaction and fertilizationin vitro in the cheetah (Acinonyx jubatus). Biol. Reprod.
46: 1047-1056.
-, Johnston, L.A., Armstrong, D.L., Simmons, L.G. and
D.E. Wildt. 1993. Birth of a Siberian tiger cub (Panthera tigris altaica) following laparoscopic intrauterine insemination. J. Zoo Wildl. Med. 24: 185- 189.
- 7Johnston, L.A., Seal, U.S., Armstrong, D.L., Tilson,
R.L., Wolf, P., Petrini, D., Simmons, L.G., Gross, T.
and D.E. Wildt. 1990. In vitro fertilization and embryo
development in vitro and in vivo in the tiger (Panthera
tigris). Biol. Reprod. 43: 733-747.

Dorst, J. and Dandelot, P. 1969. Afield guide to the larger
mammals of Africa. Collins, London.
Dotterer, M. 1992. Die Raumorganisation
von drei
benachbarten
Luchsmtinnchen(Lynxlynx) im
Schweizer Jura. [The spacing of three neighbouring
male lynx in the Swiss Jura Mountains]. Dipl. thesis,
Universityof Tubingen, Tubingen, Germany (inGerman).
Dragesco-Joffe,
A. 1993. La vie sauvage au Sahara.
[Wildlife in the Sahara.] Delachaux et Niestle,
Lausanne and Paris (in French).
Dresser, B.L., Kramer, L., Reece, B. and P.T. Russell.
1982. Induction of ovulation and successful artificial
insemination in a Persian leopard (Panthera pardus
saxicolor). Zoo Biol. 1: 55-57.
-, Gelwicks, E.J., Wachs, K.B., and G.L. Keller. 1988.
First successful transfer of cryopreserved feline (Felis
catus) embryos resulting in live offspring.
J.
Experimental 2001. 246: 180- 186.
Drucker, G.R.F. 1986. The leopards of the Bou Tferda.
Gorge. Unpubl. report, World Conservation
Monitoring Centre, Cambridge.
- . 1990. Statusof the Barbary leopard (Pantherapardus
panthera). Pp. l-4 in A. Shoemaker,ed. 1989Internatl.
leopard studbook. Riverbanks Zoological Park,
Columbia, South Carolina.
du Preez, J.S. 1970. Report on thefeeding and releaseof
30 cheetah in Etosha. Unpubl. report, Ministry of
Wildlife ConservationandTourism, EtoshaEcological
Institute, Okaukuejo, Namibia.
Dueck, H. 1990. Carnivore conservation: a proposalfor
the CanadianRockies.Unpubl. report, CanadianParks
Service, Hull, Quebec and World Wildlife FundCanada,Toronto, Ontario.
Dugan, P., ed. 1993. Wetlandsin danger: a world conservation atlas. Mitchell Beazley and IUCN, London.
Duke, K.L. 1949.Somenoteson the histology of the ovaryof the bobcat (Lynx) with special reference to thecopora lutea. Anat. Rec. 103: 11l-132.
Dunker, H. 1988.Winter studieson the lynx (Lynx lynx) insoutheasternNorway from 1960-I 982. ISSN NR 03225059, Norsk Institutt for Naturforskning, Trondheim.
Durant, S.M., Caro, T.M., Collins, D.A., Alawi, R.M. and
C.D. FitzGibbon. 1988. Migration patterns ofThomson’s gazelles and cheetahs on the Serengeti
Plains.Afr. J. Ecol. 26: 257-268.
East, R. 1992a.Conservationstatusof antelopesin North
Africa. Species18: 35-36.
- . 1992b.Conservationstatusof antelopesin Asia, part 1.
Species19: 23-25.
Easterbee,N., Hepburn, L.V. and D.J. Jefferies. 1991.
Survey of the status and distribution of the wildcat in
Scotland, 1983-l 987. Unpubl. report, Nature
ConservancyCouncil for Scotland,Peterborough.

345

References

Rev. Ecol. (Terre Vie) 43: 133-157.
- . 1990. Neotropical rainforest mammals:a field guide.
Univ. of ChicagoPress,Chicago.
- . 1991.Jaguars.Pp. 116-123 in J. Seidenstickerand S.
Lumpkin, eds.Great cats. Merehurst, London.
-, Sherman,P., Bolster, D., Goldizen, A. andJ. Terborgh.
1989. Ocelot behavior in moonlight. Pp. 233-242 in
K.H. Redford and J.F. Eisenberg, eds. Advances in
Neotropical mammalogy. Sandhill Crane Press,
Gainesville, Florida.
Enders,R.K. 1935. Mammalian life historiesfrom Barr-oColorado Island, Panama. Bull. Mus. Comp. 2002.
(Harvard) 78(4): 385-502.
Erickson, A.W. 1955. An ecological study of the bobcat
in Michigan. M.S. thesis, Mich. State Univ., East
Lansing.
Erickson, D.W. and Sampson,F.W. 1978.Impact of market dynamics on Missouri’s furbearer harvest system.
Proc. Annu. Conf Southeast.Assoc. Fish and Wildl.
Agencies32: 17-29.
Esipov, A.V. 1983. [The necessityfor the conservationof
thejungle cat.] P 58 in [Ecologicalproblemsof the conservation of wildlife.] Nauka, Moscow (in Russian).
Estes,R.D. 1967. Predatorsand scavengersI. Nat. Hist.
76(2): 20-29.
Evans, J. 1982. Plantation forestry in the tropics.
ClarendonPress,Oxford.
- . 1986. Plantation forestry in the tropics- trends andprospects.International Tree CropsJournal 4: 3- 15.
Evans, W. 1983. The cougar in New Mexico: biology, status, depredation of livestock, and managementrecommendations.Unpubl. report, New Mexico Dept. of Fish
and Game,SantaFe.
Evermann, J.F., Heeney, J.L., Roelke, M.E., McKeirnan,
A.J. and S.J. O’Brien. 1988.Biological and pathological consequencesof feline infectious peritonitis virus
infection in the cheetah.Arch. Viral. 102: 155-171.
Ewer, R.F. 1973. The carnivores. Cornell Univ. Press,
Ithaca.
Fagen,R.M. andWiley, K.S. 1978.Felid paedomorphosis,
with special reference to Leopardus. Carnivore 1( 1):72-81.
Fagotto, F. 1985.The lion in Somalia.Mammalia 49: 587588.
FAO. 1993. Summary of the final report of the Forest
ResourcesAssessment1990 for the tropical world.
Presented at the 1lth sessionof the Committee on
Forestry (8- 12 Mar 1993), United Nations Food andAgriculture Organization, Rome.
Feng, Z.-J., Cai, G.-Q. and C.-L. Zheng. 1986. [Mammalsof Tibet.] SciencePress,Beijing (in Chinese).
Fernandes,M.L. 1993. Some aspectsof the ecology andsystematicsof the wildcat (FeEissilvestris) in Portugal.
Pp. 89-93 in Proc. seminaron the biology and conser-

Eaton, R.L. 1970. Hunting behavior of the cheetah. J.
Wildl. Manage. 34( 1): 56-67.
- . 1974. The cheetah. Van Nostrand Reinhold, NewYork.
- . 1977. Reproductive biology of the leopard. 2001. Gart.
N. F. Jena 47(5): 329-35 1.
- . 1978. The conservation of the leopard in Africa:
towards an authentic philosophy of conservation.
Carnivore 1(3/4): 82- 149.
- . 1984. Survey of smaller felid breeding. 2002. Gart.
54( l/2): 101-120.
- and Verlander, K.A. 1977. Reproduction in the puma:
biology, behaviour and ontogeny. Pp. 45-70 in R.L.
Eaton, ed. The world’s cats 3: contributions to breeding, biology, behavior and husbandry. Carnivore
Research Institute, Univ. Washington, Seattle.
Edwards, S.R. and Allen, C.M. 1992. Sport hunting as asustainable use of wildlife. IUCN Sustainable Use of
Wildlife Programme, Washington, D.C.
Ehrlich, P. 1988. The loss of diversity: causes and consequences. Pp. 21-27 in E.C. Wilson and F.M. Peter, eds.
Biodiversity. National Academy Press, Washington,
D.C.
Eisenberg, J.F. 1980. The density and biomass of tropicalmammals. Pp. 35-55 in M.E. Soule and B.A. Wilcox,
eds. Conservation biology: an evolutionary-ecologicalperspective. Sinauer, Sunderland, Massachusetts.
- . 1981. The mammalian radiations: an analysis oftrends in evolution, adaptation, and behavior. Univ.
Chicago Press, Chicago.
- . 1986. Life history stategies of the Felidae: variations
on a common theme. Pp. 293-304 in S.D. Miller and
D.D. Everett, eds. Cats ofthe world: biology, conservation and management. National Wildlife Federation,
Washington, D.C.
- . 1990. Mammals c>fthe Neotropics Vol. 1: the northern
Neotropics. Univ. of Chicago Press, Chicago.
- and Lockhart, M. 1972. An ecological reconnaissanceof Wilpattu National Park, Ceylon. Smithson. Contrib.
Zool. 101: 1-118.
Ellerman, J.R. and Morrison-Scott, T.C.S. 195 1. Checklistof palaearctic and Indian mammals. British Museum of
Natural History, London.
Eloff, F.C. 1973a. Lion predation in the Kalahari Gemsbok
National Park. J. Sth. Afr. Wildl. Mgmt. Ass. 3(2): 59-63.
- . 1973b. Water use by the Kalahari lion Panthera leovernavi. Koedoe 16: 149- 154.
Elton, C: and Nicholson, M. 1942. The ten-year cycle innumbers of the lynx in Canada. J. Anim. Ecol. 11: 2 15244.
Emmons, L.H. 1987. Comparative feeding ecology of
felids in a neotropical rainforest. Behav. Ecol.
Sociobiol. 20: 27l-283.
- . 1988. A field study of ocelots(Felis pardalis) in Peru.

346

References

vation of the wildcat (Felis silvestris), Nancy, France,
23-25 September 1992. Council of Europe, Strasbourg.
Ferrari, M., Reig, O., Saltzman, J., Palermo, M.A. and B.
Marchetti. 1984. [The wild cats.] Fauna Argentina 57(in Spanish).
Ferreras, P., Aldama, J.J., Beltran, J.F., and M. Delibes.
1992. Rates and causes of mortality in a fragmented
population of Iberian lynx, Felis pardina Temminck,
1824. Biol. Conserv. 61: 197-202.
Fetisov, A.S. 1937. [Useful and harmful mammals in the
agriculture of the south Transbaikal region.] Izbjestija
Irkutsk. gos. obl. myseja, vol. 2 (in Russian).
Fey, V. 1964. The diet of leopards. African Wildlife 18:105109.
Finn, F. 1929. Sterndale ‘s mammalia of India, revised edn.
Simla, Calcutta.
Fitch-Snyder, H. In press. Environmental factors influencing captive reproduction of cheetahs. In Supplement
to the AAZPA Cheetah SSP Husbandry Protocol,
Center for Reproduction of Endangered Species,
Zoological Society of San Diego.
Fitzgerald, S. 1989. International wildlife trade: whosebusiness is it? WWF-U.S., Washington, D.C.
FitzGibbon, C.D. 1990. Why do hunting cheetahs prefermale gazelles? Anim. Behav. 40: 837-845.
Flesness, N.R. and Foose, T.J. 1990. The role of captive
breeding in the conservation of species. Pp. xi-xv in1990 IUCN red list of threatened animals. IUCN,
Gland, Switzerland and Cambridge, U.K.
Flores-Villela, 0. and Fernandez, P.G. 1989. Mexico’s
living endowment: an overview of biological diversity.
Executive summary. Conservation International andInstituto National de Investigation sobre Recursos
Bioticos (INIREB), Washington, D.C. (in English andSpanish).
Foose, T.J. and Seal, U.S. 1986. Species survival plans
for large cats in North American zoos. Pp. 173-200 in
S.D. Miller and D.D. Everett, eds. Cats of the world:
biology, conservation and management. National
Wildlife Federation, Washington, D.C.
Foreman, G.E. 1988. Behavioral and genetic analysis ofGeofroy ‘s cat (Felis geoffroyi) in captivity. Ph.D. thesis, Ohio State Univ., Columbus.
- . 1992. Pumas and people. Cat News 16: 11-12. BougyVillars, Switzerland.
~ 7 Peters, E., Somers-Griffin,
C. and K. Voigt. 1988.
Felid bibliography. Felid Research and Conservation
Interest Group and the International Society for
Endangered Cats, Columbus, Ohio.
Formozov, A.N. 1946. [Snow cover as an integral factorof the environment and its importance in the ecology
of mammals and birds.] Orig. in Russian, Engl. transl.
by the Boreal Institute, Univ. of Alberta, Edmonton.
Foster-Turley, P., MacDonald, S. and C. Mason. 1990.

Otters: an action plan for their conservation. IUCN,
Gland, Switzerland and Cambridge, U.K.
Fouraker, M. and Wildt, D. 1992. Genetic and reproductive studies of Asian lions at the Sakkarbaug Zoo and in
the Gir Forest Sanctuary. Pp. 38-40 in D.E. Wildt, J.D.
Mellen, and U.S. Seal, eds. 1992. Felid action plan,1991 and 1992: AAZPA Felid Taxon Advisory Groupregional collection plan and IUCN Captive BreedingSpecialist Group global felid action plan. National
Zoological Park, Front Royal, Virginia.
Fowler, M.E. 1986. Felidae. Pp. 831-841 in M.E. Fowler,
ed. Zoo and wild animal medicine, 2d edn. W.B.
Saunders, Philadelphia.
Fox, J.F. 1978. Forest fires and the snowshoe hare-Canada
lynx cycle. Oecologia 3 1: 349-374.
Fox, J.L. 1985. An observation of lynx in Nepal. J.
Bombay Nat. Hist. Sot. 82: 394.
- . 1989. A review of the status and ecology of the snow
leopard (Panthera uncia). International Snow Leopard
Trust, Seattle.
- . 1994. Snow leopard conservation in the wild-a comprehensive perspective on a low density and highly
fragmented population. In J.L. Fox and Du Jizeng, eds.
Proc. seventh international snow leopard symposium.
International Snow Leopard Trust, Seattle.
- and Chundawat, R.S. 1988. Observations of snow leopard stalking, killing, and feeding behavior. Mammalia
52(l): 137-140.
- and Du Jizeng, eds. 1994. Proc. of the seventh international snow leopard symposium. International Snow
Leopard Trust, Seattle.
-, Nurbu, C. and R.S. Chundawat. 1991 b. The mountain
ungulates of Ladakh, India. Biol. Conserv. 58: 167- 190.
-, Sinha, S.P., Chundawat, R.S. and P.K. Das. 1991a.
Status of the snow leopard Panthera uncia in
Northwest India. Biol. Conserv. 55: 283-298.
Frame, G.W. 1977. Cheetah ecology and behaviour. Pp.
74-87 in 1976-76 Annual Report, Serengeti ResearchInstitute, Arusha.
- . 1980. Cheetah social organization in the Serengeti
ecosystem, Tanzania. Paper presented at the meetingof the Animal Behavior Society, Fort Collins,
Colorado.
- . 1984. Cheetah. Pp. 40-43 in D. Macdonald, ed. The
encyclopedia of mammals. Facts on File, New York.
- . 1992. First record of the king cheetah in West Africa.
Cat News 17: 2-3. Bougy-Villars, Switzerland.
- and Frame, L.H. 1980. Cheetahs: in a race for survival.
Nat. Geog. May: 712-728.
- . 198 1. SwifiL and enduring: cheetahs and wild dogs of
the Serengeti. E.P. Dutton, New York.
- . 1984. Cheetah male cooperation: test of a mutualism
model. Paper presented at the meeting of the Animal
Behavior Society, Cheney, Washington.

347

References

Manage. 23: 226-229.
Gasperetti, J., Harrison, D.L., and W. Biittiker. 1986. The
Carnivora of Arabia. Fauna of SaudiArabia 7: 397-461.
Gaumer, G.F. 1917. [Monograph on mammals of
Yucatan.] Dept. Talleres Graficos, Secretaria de
Fomento, Mexico (in Spanish).
Gee, E.P. 1964.The wildltfe of India. Collins, London.
Geertsema,A.A. 1976. Impressionsand observationson
serval behaviour in Tanzania, East Africa. Mammalia
40(l): 13-19.
-. 1985.Aspectsof the ecology of the servalLeptailurus
serval in the NgorongoroCrater, Tanzania.Netherlands
J. 2001.35(4): 527-610.
Genovesi, P. and Boitani, L. 1993. Spacing patterns andactivity rhythms of a wildcat (Felis silvestris) in Italy.
Pp. 9% 101 in Proc. seminaron the biology and conservation of the wildcat (Felis silvestris), Nancy,
France, 23-25 September1992. Strasbourg: Councilof Europe.
The Gentleman’s Magazine. 1793. Reproduced in N.
Courtney. 1980. Tiger: symbol of freedom. Quartet,
London.
Ghosh,A. 1988.Freedomagain. TelegraphSundaySuppl.
23 July, Calcutta, India.
-. 1994.Alarming decline in number of tigers. Timesof
India, 9 May, New Delhi.
Gibb, J.A. 1990.The Europeanrabbit. Pp. 116-120in J.A.
Chapman and J.E.C. Flux, eds. Rabbits, hares andpikas: status survey and conservation action plan.
IUCN Gland, Switzerland and Cambridge,U.K.
Gieteling, C. 1972.Jaguar en ocelot: biologies,bedreiging
en bescherming. [Jaguar and ocelot: biology, threatsand protection.] Unpubl. report, WWF Netherlands,
AA Zeist (in Dutch with English abstract).
Gilbert, D.A., Packer, C., Pusey, A.E., Stephens,J.C. and
S.J. O’Brien. 1991. Analytical DNA fingerprinting inlions: parentage, gen.etic diversity, and kinship. J.
Hered. 82: 378-386.
Gilpin, M.E. 1973.Do hareseat lynx? Am. Nat. 107: 727730.
- and Soul& M.E. 1986. Minimum viable populations:processesof speciesextinction. Pp. 19-34in M. Soul&
ed. Conservation biology: the scienceof scarcity anddiversity. Sinaeur,Sunderland,Massachusetts.
Gipps, J. 1991.Preface in Beyond captive breeding: reintroducing endangered mammalsto the wild. Oxford
Scientific Publ., Oxford.
Ginsberg, J.R. and MacDonald, D.W. 1990. Foxes,
wolves,jackals and dogs: an action plan for the conservation of canids. IUCN, Gland, Switzerland andCambridge,U.K.
Gittleman, J.L. 1985.Carnivore body size: ecological andtaxonomic correlates.Oecologia 67: 540-554.
Glade, A., ed. 1988.Red list of Chilean terrestrial verte-

Fredrickson, L.F. and Rice, L.A. 1979. Bobcat management survey study in South Dakota, 1977- 1979. Proc.
bobcat res. con&, natl. wildl. fed. sci. tech. ser. 6: 32-36.
Freeman, H. 1975. A preliminary study of the behaviour ofcaptive snow leopards. Int. Zoo Yearb. 15: 217-222.
-. 1977. Social behavior in the snow leopard. Pp. 94-98 in
American Associationof Zoological Parks andAquariums Annual Proceedings. San Diego.
- 9ed. 1988. Proc. ofthefifih international snow leopard
symposium. International Snow Leopard Trust andWildlife Institute of India. Conway, Bombay.
Fritts, S.H. 1973. Age, food habits, and reproduction of thebobcat (Lynx rufus) in Arkansas. M.S. thesis, Univ.
Arkansas, Fayetteville.
- and Sealander, J.A. 1978a. Diets of bobcats in Arkansas
with special reference to age and sex differences. J.
Wildl. Manage. 42: 533-539.
-. 1978b. Reproductive biology and population characteristics of bobcats (Lynx rufis) in Arkansas. J. Mamm.
59: 347-353.
Fuller, K.S., Swift, B,. Jorgensen, A., and A. Brautigam.
1987. Latin American wildlife trade laws, 2d edn.
(rev.). WWF-U.S., Washington, D.C.
Fuller, T.K., Berg, W.E., and D.W. Kuehn. 1985. Survival
rates and mortality factors of adult bobcats in northcentral Minnesota. J. Wildl. Manage. 49: 292-296.
-, Biknevicius, A.R. and P.W. Kat. 1988. Home rangeof an African wildcat, Felis silvestris (Schreber) near
Elmenteita, Kenya. 2. Saugetierk. 53: 380-38 1.
Gaillard. 1969. [On the presence of the golden cat (Felis
aurata Temm.) and caracal (Felis caracal Schreb.) insouthern Senegal.] Mammalia 33: 350-35 1 (in French).
Gao, Yaoting et al. 1987. [Fauna Sinica. Mammalia, vol.
8: Carnivora.] Science Press, Beijing (in Chinese).
Garcia-Perea, R. 1992. New data on the systematics
of lynxes. Cat News 16: 15-16. Bougy-Villars,
Switzerland.
-. 1994. The pampas cat group (Genus Lynchailurus
Severtzov, 1858) (Carnivora, Felidae): a systematic andbiogeographic review. Amer. Mus. Novit.
Gardner, A.L. 1971. Notes on the little spotted cat, Felis
tigrina oncilla Thomas, in Costa Rica. J. Mammal.
52(2): 464-465.
Garga, D.P. 1948. How far can a tiger swim? J. Bombay
Nat. Hist. Sot. 47(3): 545.
Garland, T., Jr. 1983. The relation between maximal running speed and body mass in terrestrial mammals. J.
Zool., Lond. 199: 157-170.
Garshelis, D.L. 1992. Mark-recapture density estimation
for animals with large home ranges. Pp. 1098-l 111 in
D.R. McCullough and R.H. Barrett, eds. WiZdEife 2001:
populations. Elsevier, London.
Gashwiler, J.S., Robinette, W.L. and O.W. Morris. 1960.
Foods of bobcats in Utah and eastern Nevada. J. Wildl.

348

References

Unpubl. revised status report for the Committee on
International Trade in EndangeredSpecies.
Govt. of China. 1994. Proposalto register the first commercial captive-breeding operation for Siberian tiger.
Proposalsubmittedto the CITES Secretariat,Lausanne,
Switzerland.
Govt. of India. 1988. The snow leopard conservation
scheme. Ministry of Environment and Forests,
Government of India, New Delhi.
- . 1991. The stateofforest report, 1991.Forest Survey ofIndia, Ministry of Environment and Forests, Dehra
Dun.
- . 1993. A review of Project Tiger. Ministry ofEnvironment and Forests,Government of India, New
Delhi.
Govt. of Namibia. 1992. Proposal to transfer Acinonyx
jubatus (populations of Botswana, Malawi, Namibia,
Zambia, Zimbabwe) from Appendix I to Appendix II.
CITES Ref Dot. 8.46 no. 9. CITES Secretariat,
Lausanne,Switzerland.
Govt. of U.S. 1983a.Proposalto remove Lynx rufis (populations of the United States and Canada) from
Appendix II. Proc. Conf of the Parties to CITES 4,
CITES Secretariat,Lausanne,Switzerland.
- . 1983b.Proposalto remove Lynx canadensis(populations of the United Statesand Canada)from Appendix
II. Proc. Conf of the Parties to CITES 4, CITES
Secretariat,Lausanne,Switzerland.
- . 1992.Proposalto transfer FeZisrufa escuinapaefrom
Appendix I to Appendix II. Proc. Conf of the Parties to
CITES 8. CITES Secretariat,Lausanne,Switzerland.
Graells, M.P. 1897. [Iberian mammalfauna.] Mem. Real
Acad. Ciencias17: 224-229.Madrid (in Spanish).
Graham, A. 1966.EastAfrican Wild Life Society cheetah
survey: extracts from the report by wildlife services.E.
Afr. Wildl. J. 4: 50-55.
Gray, J.E. 1867.Notes on the skulls of the cats (Felidae).
Proc. 2001.Sot. Lond. 1867: 258-277, 874-876.
Green, M.J.B. 1987. The conservation status of the leopard, goral and serow in Bangladesh,Bhutan, northern
India and southern Tibet. Unpubl. report, World
ConservationMonitoring Centre, Cambridge.
- . 1988. Protected areasand snow leopards:their status.
Pp. 3-10 in H. Freeman, ed. Proc. fifth international
snowleopard symposium.International Snow Leopard
Trust andWildlife Institute of India. Conway, Bombay.
- 7camp. 1993. Nature reserves of the Himalaya andmountainsof Central Asia. Oxford Univ. Press,New
Delhi for IUCN.
- . 1994. Protecting the mountains of Central Asia andtheir snow leopard populations. In J.L. Fox and Du
Jizeng, eds.Proc. seventhinternational snow leopard
symposium.International Snow LeopardTrust, Seattle.
Green, R. 1991. Wild cat speciesof the world. Basset,

brates. Chilean Forest Service (CONAF), Santiago.
Glass, G. and Todd, N. 1977. Quasi-continuous variationof the second upper premolar in FeZis bengalensis Kerr,
1972 and its significance for some fossil lynxes.
Sonderdr. 2. Saugetierk. 42: 36-44.
Goebel, A.M. and Whitmore, D.H. 1987. Use of electrophoretic data in the reevaluation of tiger systematics.
Pp. 36-50 in R.L. Tilson and U.S. Seal, eds. Tigers of
the world: the biology, biopolitics, management, andconservation of an endangered species. Noyes, Park
Ridge, New Jersey.
Goldman, D. and O’Brien, S.J. 1993. Comparative studies of proteins employing two dimensional electrophoresis. Meth. Enzymol.
Goldman, E.A. 1920. Mammals of Panama. Smithsonian
Misc. Coil. 69(5): l-309.
Gonyea, W.J. 1976. Adaptive differences in the body proportions of large felids. Acta. Anat. 96: 8 l-96.
Gonzlez, C.B. and Leal, C.G. 1984. [Forest mammals of
the Mexican Basin.] Programme on Man and the
Biosphere (UNESCO) and Editorial Limusa, Mexico
City (in Spanish).
Goodman, S.M. and Helmy, I. 1986. The sand cat Felis
margarita Loche, 1858 in Egypt. Mammalia 50: 120123.
Goodrowe, K.L., Wall, R.J., O’Brien, S.J., Schmidt, P.M.
and D.E. Wildt. 1988. Developmental competence ofdomestic cat follicular oocytes after fertilization invitro. Biol. Reprod. 39: 355-372.
Goodwyn, F., Jr. 1970. Behavior, life
. history and present
status of the jaguarundi, Felis yagouaroundi(Lacepede) in south Texas. M.A. thesis, Texas A & I
Univ., Kingsville.
Gossow,H. and Honsig-Erlenburg,P. 1986.Management
problemswith reintroduced lynx in Austria. Pp. 77-83
in S.D. Miller and D.D. Everett, eds.Catsofthe world:
biology, conservation and management. National
Wildlife Federation,Washington, D.C.
Gould, G.I. 1989. Bobcat harvest assessment,1987-88.
Technical report 1989-3, NongameBird and Mammal
Section report, Dept. of Fish and Game, Sacramento,
California.
Gouttenoire, G. 1954. [Hunting in Tunisia]. In [The big
book of the fauna of Africa and its hunting.] Geneva(in French).
Govt. of Brazil. 1992.Proposalto transfer FeZisgeofSroyi
from Appendix II to Appendix I. Proc. Conf of theParties to CITES 8. CITES Secretariat, Lausanne,
Switzerland.
Govt. of Canada. 1983. Proposal for the deletion of theCanadianpopulation of the bobcat, Lynx rufus, from
Appendix II. Proc. Conf of the Parties to CITES 4,
CITES Secretariat,Lausanne,Switzerland.
- . 1988.Statusof the lynx (Lynx canadensis)in Canada.

349

References

atics of Old World Felidae. Carnivore 5(2): 35-46.
- . 1990.Report on project no. IRA/87/014/A/OI/12: visit
to protected areas in Iran. Unpubl. report, United
NationsDevelopment Programme.
Grzimek, B. 1975. Grzimek ‘sAnimal Life Encyclopaedia
12: 354. Van NostrandReinhold Co. New York.
Gudynas, E. 1989. The conservation status of SouthAmerican rodents: many questionsbut few answers.
Pp. 20-25 in W.Z. Lidicker, Jr., ed. Rodents: A world
survey of species of conservation concern. IUCN
Species Survival Comm. Oct. Paper no. 4, Gland,
Switzerland.
Guggisberg, C.A.W. 1961. Simba: the life of the lion.
Howard Timmins, CapeTown.
- . 1975. Wild cats of the world. David and Charles,
London.
Gui Xiaojie and Meng Sha. 1993. The challenge andstrategiesfor managementof the South China tiger
Panthera tigris amoyensis. Paper presented at the
Global Tiger Forum, 4 March 1994,New Delhi, India.
Gyldenstolpe, N. 1928.Zoological resultsof the Swedish
expedition to central Africa 1921, V: Mammals fromthe Virunga Volcanoes,north of Lake Kivu. Ark. 2001.)
Stockholm 20A (4): l-76.
Habibi, K. 1977. The mammalsof Afghanistan: their distribution and status.Unpubl. report to the UNDP, FAO
andMinistry of Agriculture, Kabul.
Habijan,V. and Dimitrijevic, S. 1979.Feedingof the wildcat, Felis silvestrisSchreber1777,in Vojvodina. Drugi
KongresEkologia Jogoslavie (Zagreb): 1447-1456.
Haglund, B. 1966. [Winter habitsof the lynx and wolverine asrevealed by tracking in snow.] SwedishWildlife
4: 81-299 (in Swedish).
Hall, E.R. 1981.Mammalsof North America, 2d edn.John
Wiley, New York.
- and Dalquest,W.W. 1963.The mammalsof Veracruz.
Univ. KansasPubl., Mus. Nat. Hist. 14: 165-362.
Hall, H.T. andNewsom,J.D. 1976.Summerhomeranges
andmovementsof bobcatsin bottomlandhardwoodsof
southern Louisiana. Proc. Annu. Conf. Southeast.
Assoc.Fish and Wildl. Agencies30: 422-436.
Haller, H. 1992. Zur okologie desLuchsesLynx lvnx im
Verlauf seiner Wiederansiedlung in den Walliser
Alpen. [Ecology of the lynx (Lynx lynx) during its reintroduction in the Valais, Swiss Alps.] Mammalia
depicta 1.5,2. Saugetierk.Paul Parey, Hamburg.
Haller, H. and Breitenmoser, U. 1986. Zur Raumorganisationder in den Schweizer Alpen wiederangesiedelten Population desLuchsesLynx lynx. [Spatialorganization of a reintroduced lynx population in the
Swiss Alps.] 2. Saugetierk. 51(5): 289-3 11 (inGerman).
Halloy, S. 1985. Climatologia y edafologia de alta montafia en relation con la composition y adaptation de

Plymouth.
Greer, J.K. 1965a. Mammals of Malleco Province, Chile.
Pub. Mich. St. Univ. Mus.: Biol. Ser. 3: 49-l 52.
- . 1965b. Another record of the Andean Highland Catfrom Chile. J. Mammal. 46: 507.
Griffith, M.A. and Fendley, T.T. 1986. Pre- and post-disperal movement behavior of subadult bobcats on the
Savannah River Plant. Pp. 277-290 in S.D. Miller and
D.D. Everett, eds. Cats ofthe world: biology, conservation and management. National Wildlife Federation,
Washington, D.C.
Griffiths, M. 1993. Management of large mammals.
Unpubl. project report, WWF International, Gland,
Switzerland.
- and van Schaik, C.P. 1993a. The impact of humanactivity on the abundance and activity periods ofSumatran rain forest wildlife. In Griffiths, M. 1993.
Management of large mammals. Unpubl. report, WWF
International, Gland, Switzerland.
- . 1993b. Camera-trapping: a new method to study elusive rain forest animals and a tool for conservation education. In Griffiths, M. 1993. Management of large
mammals. Unpubl. report, WWF International, Gland,
Switzerland.
Grimwood, I.R. 1969. Notes ofthe distribution and status
of some Peruvian mammals in 1968. Special publ. no.
2 1, New York Zoological Society, Bronx.
Grisham, J. 1988. AAZPA Species Survival Plan: Cheetah
husbandry and reproductive survey and guidelines.
Oklahoma City Zoological Park, Oklahoma City.
Grobler, J.H. 198 1. Feeding behaviour of the caracal Felis
caracal Schreber, 1776, in the Mountain Zebra
National Park. S. Afr. J. 2001. 16: 259-262.
~ and Wilson, V.J. 1972. Food of the leopard Panthera
pardus (Linn.) in the Rhodes Matopos National Park,
Rhodesia, as determined by faecal analysis. Arnoldia
5(35): l-10.
Groombridge, B. 1988. Baluchistan province, Pakistan: apreliminary environmental profile. Unpubl. report,
World Conservation Monitoring Centre, Cambridge.
- 9ed. 1993. 1994 IUCN Red List of threatenedanimals.
IUCN, Gland, Switzerland andCambridge,U.K.
Gros, P. 1990. Global cheetahproject PhaseI: Cheetah
status in southern Africa. Unpubl. report., Univ. of
California, Davis.
Grosjean, D. 1992. The impact of lynx on livestock. Pp.
66-70 in The situation, conservation needsand reintroduction of lynx in Europe. Proc. symp. 17-I 9
October, Neuchatel. Council of Europe, Strasbourg.
Grove, N. 1981. Wild cargo: the businessof smuggling
animals.Nat. Geog. 159(3): 287-316.
Groves, C.P. 1980.The Chinesemountaincat (Felis bieti).
Carnivore 3(3): 35-41.
- . 1982.Cranial anddental characteristicsin the system-

350

References

las comunidades bioticas (con especial referencia a las
Cumbres Calchaquies, Tucuman. [Climatology andedaphology of high mountains in relation to the composition and adaptation of biotic communities (withspecial reference to the Cumbres Calchaquies, Tucuman)]. Ph.D. thesis. Fat. Cs. Nat. UNT. University
Microfilms Int. publ. #85-02967. Ann Arbor, Michigan.
Haltenorth, T. 1953. Lebende arabische Sandkatze (F&smargarita Loche, 1858). [Living Arabian sandcats.]
Saugetierk. Mitt. 1: 7 l-73.
-. 1957. [The wildcat.] Die neue Brehm Bucherei,
Wittenberg-Lutherstadt.
Hamer, M. 1992. Poachers kill tigers for their bones. New
Scientist 11 July: 5.
Hamilton, D.A. 1982. Ecology ofthe bobcat in Missouri.
M.S. thesis, Univ. of Missouri, Columbia.
Hamilton, P.H. 1976. The movements ofleopards in Tsavo
National Park, Kenya, as determined by radio-tracking. M.S. thesis, Univ. of Nairobi, Nairobi.
-. 198 1. The leopard (Panthera pardus) and the cheetah(Acinonyx jubatus) in Kenya. Unpubl. report to the
U.S. Fish and Wildlife Service.
-. 1986a. Status of the cheetah in Kenya, with reference
to sub-Sah&an Africa. Pp. 65-76 in S.D. Miller and
D.D. Everett, eds. Cats ofthe world: biology, conservation and management. National Wildlife Federation,
Washington, D.C.
-. 1986b. Status of the leopard in Kenya, with reference
to sub-Saharan Africa. Pp. 447-459 in S.D. Miller and
D.D. Everett, eds. Cats of. the world: biology, conservation and management.National Wildlife Federation,
Washington,D.C.
Hanby, J.P. and Bygott, J.D. 1979.Population changesin
lions and other predators. Pp. 249-262 in A.R.E.
Sinclair and M. Norton-Griffiths, eds. Serengeti:
Dynamics ofan ecosystem.Chicago: Univ. Chicago
Press.
- . 1987.Emigration of subadultlions. Anim. Behav. 35:
161-169.
-. 1991. Lions. Pp. 80-93 in J. Seidensticker and S.
Lumpkin, eds.Great cats. Merehurst, London.
Hansen,K. 1992. Cougar: the American lion. Mountain
Lion Foundation and Northland Press, Flagstaff,
Arizona.
Hanski,I., Hansson,L., andH. Henttonen. 1991.Specialist
predators,generalistpredators,andthe microtine rodent
cycle. J. Anim. Ecol. 11: 2 15-244.
Happold, D.C.D. 1984. Small mammals.Pp. 25 l-275 in
J.L. Cloudsley-Thompson, ed. Sahara Desert.
PergamonPress,Oxford.
- . 1987. The mammalsof Nigeria. Oxford Univ. Press,
Oxford.
Haque, M.N. 1988. Scavenging habit of fishing cat. J.
Bombay.Nat. Hist. Sot. 85(l): 183.

Hardy, E. 1947. The Palestine leopard. J. Sot. Preserv.
Fau. Emp., y2.s.55: 16-20.
Hardy, I.W. 1979. Golden cat in the Aberdare National
Park. East Afr. Nat. Hist. Sot. Bull. 1979: 11l-l 12.
Harestad,A.S. and Bunnell, F.L. 1979. Home range andbody weight-a reevaluation.Ecol. 60: 389-402.
Harpster, J. 1990. Floridians fight to save panthers.
Christian ScienceMonitor, 24 April.
Harrington, F.A., Jr. 1977. A guide to the mammalsof
Iran. Dept. of the Environment, Govt. of Iran, Tehran.
Harris, R.B. and Allendorf, F.W. 1989.Genetically effective population sizesof large mammals:an assessmentof indicators. Conserv.Biol. 3(2): 181- 191.
Harrison, D.L. 1983. The mammalfauna of Oman withspecialreference to conservation and the Oman FloraandFauna Survey. J. Oman Studies6: 329-339.
- andBates,P.J.J. 1991.The mammalsofArabia, 2d edn.
HarrisonZoological Museum, Sevenoaks,U.K.
Harrison, J. 1974. An introduction to mammals ofSingapore and Malaya. Malayan Nature Society,Singapore.
Hart, J.A. and Katembo, M. In prep. Diet and preyd selection in an African,forest,felid community.
Hassinger, J. D. 1965. A survey of the mammals of
Afghanistan: results of the 1965 Street Expedition.
Fieldiana 2001.60: 1- 196.
Hast, M.H. 1989. The larynx of roaring and non-roaring
cats.J. Anat. 163: 117-121.
Hatt, R.T. 1959. The mammals of Iraq. Museum of
Zoology, Univ. of Michigan, Ann Arbor.
Headley, K. 1992. Trade in exceptional circumstances:an analysisof commercein Appendix I animal species.
Unpubl. report, TRAFFIC International, Cambridge.
Heaney, G.F. 1943. Occurrence of lions in Persia. J.
BombayNat. Hist. Sot. 44: 467.
Hecketsweiler,P.L. 1988. [Conservationand rational use
offorest ecosystemsin central Africa: national reportfor the Congo.] Unpubl. report, IUCN, Gland,
Switzerland (in French).
Heeney, J., Evermann, J., McKeiman, A., Marker-Kraus,
L., Roelke,M., Bush, M., Wildt, D., Meltzer, D., Colly,
L., Lukas, J., Manton, V., Caro, T., and S. O’Brien.
1990.Prevalenceandimplicationsof feline coronavirus
infections of captive and free-ranging cheetahs(Acinonyx jubatus). J. Virol. 64(5): 1964-1972.
Hell, P. 1973.’(Resultsof lynx researchin Czechoslovakia.
Part I: Nutritional ecology and importanceto hunting.]
Beitr zur Jagd- und Wildforsch 8: 335-344 (inGerman).
- . 1992. Managing the lynx population in Czechoslovakia. Pp. 33-35 in The situation, conservation needs
and reintroduction of lynx in Europe. Proc. symp. 1719 October, Neuchatel. Council of Europe, Strasbourg.
Hemker, T.P. 1982.Population characteristicsand move-351

References

Cambridge.
Grubb, P. and C.P. Groves. 1976. Notes on the sandcat, Felis margarita Loche, 1858. Z. Saugetierke. 41:
286-303.
- and Schutt, G. 1973. [Pleistocene leopards (Panthera
pardus) of Java and southern China.] Koninkl. Neder.
Akad. Wetensch. Proc. Sers. B 76: 37-49 (in German).
Hendrichs, H. 1975. The status of the tiger in the
Sundarbans mangrove forest (Bay of Bengal).
Saugetierk. Mitt. 3 : 16 1- 199.
Heptner, V.H. 1970. [The Turkestan sand dune cat(Barchan cat), Felis margarita thinobia Ogn., 1926.1 D.
Zool. Gart. 39: 116-128 (in German).
-. 197 1. [On the systematic position of the Amur forest
cat and some other east Asian cats placed in Felis bengalensis Kerr, 1792.1 Zool. Zhurn. 5 l( 11) (in Russian).
- and Sludskii, A.A. 1972. [Mammals of the Soviet
Union. Vol III: Carnivores (Feloidea).] Vyssha Shkola,
Moscow (in Russian). Engl. transl. edited by R.S.
Hoffmann, Smithsonian Inst. and the Natl. Science
Fndn., Washington DC, 1992.
Herrenschmidt, V. and Leger, F. 1987. [The lynx Lynx
Lynx in north-eastern France: colonization of the French
Jura mountains and reintroduction of the species in the
Vosges mountains.] Ciconia 1l(2): 13 l-5 1 (in French).
- and Vandel, J.M. 1989. Premier bilan des degats occasiones par les lynx sur les troupeaux d ‘animaux domestiques de la chaine du Jura. [A preliminary assessmentof the damage caused by lynx to domestic animals in
the Jura mountains.] Unpubl. report, Office National dela Chasse, Station d’Alsace et du Massif Vosgien.
Herrington, S.J. 1986. Phylogenetic relationships of thewild cats of the world. Ph.D. thesis, Univ. Kansas,
Lawrence.
-. 1987. Subspecies and the conservation of Panthera
tigris: preserving genetic heterogeneity. Pp. 5 l-63 in
R.L. Tilson and U.S. Seal, eds. Tigers of the world: thebiology, biopolitics, management, and conservation ofan endangered species. Noyes, Park Ridge, New
Jersey.
Hewson, R. 1983. The food of wild cats (Felis silvestris)
and red foxes (Vulpes vulpes) in west and northeast
Scotland. J. Zool., Lond. 200: 283-289.
Heywood, V.H. and Stuart, S.N. 1992. Species extinctions
in tropical forests. Pp. 91-l 18 in T.C. Whitmore and
J.A. Sayer, eds. Tropical deforestation and species
extinction. Chapman and Hall, London.
Hildebrand, M. 1959. Motions of the running cheetah andhorse. J. Mammal. 40: 48 l-495.
-. 196 1. Further studies on locomotion of the cheetah. J.
Mammal. 42: 84-91.
Hillard, D. 1989. Vanishing tracks: four w
years among thesnow leopards of Nepal. William and Morrow, NewYork.

mentpatterns of cougars in southern Utah. M.S. thesis, Utah State Univ., Logan.
Hemker, T.P., Lindzey, F.G., Ackerman, B.B. and A.J.
Button. 1984. Population characteristics and movement
patterns of cougars in southern Utah. J. Wildl. Manage.
48(4): 1275 1284.
-. 1986. Survival of cougar cubs in a non-hunted population. Pp. 327-332 in S.D. Miller and D.D. Everett,eds. Cats of the world: biology, conservation and management. National Wildlife Federation, Washington,
D.C.
Hemmer, H. 1968. Untersuchungenzur Stammesgeschichte der Panterkatzen (Pantherinae) II: Studien
zur Ethologie des Nebelparders Neofelis nebulosa(Griffith 1821) und des Irbis Uncia uncia (Schreber1775). [Studies of the phylogenetic history of thePantherinae II: research into the ethology of the clouded
leopard and the snow leopard.] Veroff Zool.
Staatssamml. Munchen 12: 155-247 (in German).
-. 1972. Uncia uncia. Mammal. Species 20: l-5.
-. 1974a. [Studies on the systematics and biology of thesand cat.] Zt. d. Kiilner Zoo 17( 1): 11-20 (in German).
-. 1974b. [Studies of the phylogenetic history of the pantherines (Pantherinae) III. On the races of the lion
Panther-a leo (Linnaeus,
1758).] Veroff. 2001.
Staatssamml. Munchen 17: 167-280.
-. 1976. Gestation period and postnatal development infelids. Pp. 143-165 in R.L. Eaton, ed. The world’s cats
3. Carnivore Research Institute, Univ. Washington,
Seattle.
-. 1977. Biology and breeding of the sand cat. Pp. 13-20
in R.L. Eaton, ed. The world’s cats 3: contributions tobreeding, biology, behavior and husbandry. Carnivore
Research Institute, Univ. Washington, Seattle.
-. 1978a. The evolutionary systematics of living Felidae:
present status and current problems. Carnivore l( 1):7 l-79.
-.
1978b. Zur intraspezifischen,geographischen
Variabilitat des Tigers (Panthera tigris L.) nebst
Anmerkungen zu taxonomischen Fragen. [On theintraspecific and geographic variability of the tiger withremarks on taxonomic questions.] Pp. 60-64 in[Congress rept. first int. symp. mgt. & breeding of thetiger.] Int. tiger studbook, Leipzig Zool. Garten,
Leipzig.
- * 1987. The phylogeny of the tiger (Panthera tigris).
Pp. 28-35 in R.L. Tilson and U.S. Seal, eds. Tigers of
the world: the biology, biopolitics, management, andconservation of an endangered species. Noyes, Park
Ridge, New Jersey.
-. 1988. Asiatic cheetah. Cat News 9: 13. Bougy-Villars,
Switzerland.
-. 1990. Domestication: the decline of environmental
appreciation,
2d edn. Cambridge Univ. Press,

-,

352

References

Hills, D.M. and Smithers, R.H.N. 1980. The “King cheetah”: a historical review. Amoldia 9( 1): l-23.
Hjelm, K. 1991. [Lynx status, Jamtlands county, 1991.1
Unpubl. report,
SwedishSociety for Nature
Conservation, Stockholm (in Swedish).
Holdridge, L.R. 1978. Resource use and conservation in
the high mountain Andes. Pp. 57-68 in The use of highmountains of the world. Department of Lands andSurvey and IUCN, Wellington, New Zealand.
Hoogesteijn, R., Hoogesteijn, A. and E. Mondolfi. 1993.
Jaguar predation vs. conservation: cattle mortality by
felines on three ranches in the Venzuelan llanos. In N.
Dunstone and M.L. Gorman, eds. Mammals as predators. Proc. Symp. 2001. Sot. Lond. 65. Clarendon,
Oxford.
- and Mondolfi, E. 1992. The jaguar. Ediciones
Armitano, Caracas, Venezuela (in Spanish, English,
German, French, Italian).
Hoogerwerf, A. 1970. Udjung Kulon: the land of the last
Javan rhinoceros. E. J. Brill, Leiden.
Hopkins, R.A. 1989. Ecology of the puma in the Diablo
Range, Cabfornia. Ph.D. thesis, Univ. of California,
Berkeley.
Hoppe-Dominik, B. 1984. [Study of the prey range of theleopard, Panthera pardus, in the Tai National Park,
Ivory Coast.] Mammalia 48(4): 477-487 (in French).
Hopwood, A.T. 1947. Contributions to the study of some
African mammals. III: Adaptation in the bones of thefore limb of the lion, leopard and cheetah. J. Linn. Sot.
2001. 41: 259-27 1.
Hornocker, M.G. 1969. Winter territoriality in mountainlions. J. Wildl. Manage. 33(3): 457-464.
- . 1970. An analysis of mountain lion predation uponmule deer and elk in the Idaho Primitive Area. Wildl.
Monogr. 21: l-39.
- . 1992. Learning to live with mountain lions. Nat. Geog.
182( 1): 52-65.
Hose, C. 1893. Mammals ofBorneo. British Museum Nat.
Hist., London.
Housse, P.R. 1953. [Wild animals of Chile.] Ediciones
Universidad de Chile, Santiago(in Spanish).
Howard, J.G., Besour, M.A., Donoghue, A.M. and D.E.
Wildt. 1992b.The effect of pre-ovulatory anaesthesia
on ovulation in laparascopicallyinseminateddomestic
cats.J. Reproductionand Fertility 96: 175-186.
- 7 Donoghue, A.M., Goodrowe, K.L., Blumer, E.,
Snodgrass,K., Starnes,D., Tucker, M., Bush, M. and
D.E. Wild. 1992a. Successful induction of ovarian
activity and laparoscopicintrauterine artificial insemination in the cheetah(Acinonyx jubatus). J. Zoo Wildl.
Med. 23: 288-300.
Hu Jinzhu and Wang Youzi, eds. 1984. [Sichuan Fauna
Economica.Vol. 2: Mammals.] Chengdu(in Chinese).
Hubbard, A.L., McOrist, S., Jones,T.W., Boid, R., Scott,353

R. andN. Easterbee.1992.Is survival of Europeanwildcats Fells silvestrisin Britain threatenedby interbreeding with domesticcats?Biol. Conserv.61: 203-208.
Hucht-Ciorga, I. 1988. [Studies in lynx biology: hunting
behavior, utilization of carcasses,interspecific communication, andphysical characteristicsthat canbe recognized in tracks.] Schriften des Arbeitskreises
Wildbiologie und Jagdwissenschaft an der JustusLiebig- Universitat Giessen 19. Ferdinand Enke,
Stuttgart (in German).
Hufnagl, E. 1972.Libyan mammals.OleanderPress,U.K.
Hulley, J.T. 1976. Maintenance and breeding of captive
jaguarundis at Chester Zoo and Toronto. Intl. Zoo
Yearb. 16: 120-122.
Hummel, M. 1990.A conservationstrategyfor large carnivores in Canada. World Wildlife Fund-Canada,
Toronto, Ontario.
Humphrey, S.R. andBain, J.R. 1990.Endangeredanimalsof Thailand. SandhillCranePress,Gainesville,Florida.
- and Foster, M.L. 1991. EfSectivenessof wildlife crossing structures on Alligator Alley (I-75) for reducing
animal/auto collisions. Annual performance report, 1
July 1990-30 June 1991, Florida Game and Fresh
Water Fish Commission,Tallahassee.
Husain, K.Z. 1974. An introduction to the wildlife of
Bangladesh.F. Ahamed, Dacca.
Husson,A.M. 1978. The mammalsof Surinam. EJ Brill,
Leiden.
Hutchins, M. and Wiese, R.J. 1991. Beyond genetic anddemographic management:the future of the Species
Survival Plan andrelatedAAZPA conservationefforts.
Zoo Biol. 10: 285-292.
- 7Foose,T. and U.S. Seal. 1991. The role of veterinary
medicine in endangeredspeciesconservation. J. Zoo
Wildl. Med. 22(3): 277-281.
Ilani, G. 1990. Leopard Panthera pardus in Israel. Cat
News 12: 4-5. Bougy-Villars, Switzerland.
- . 1981. The leopardsof the Judeandesert.Israel Land &Nature 6(2): 59-71.
Imaizumi, Y. 1967.A new genusand speciesof cat from
Iriomote, Ryuku Islands.J. Mammal. Sot. Japan 3: 75105.
Inoue, T. 1972.The food habits of the Tsushimaleopard
cat, Felis bengalensisssp.,analyzed from their scats.
J. Mammal. Sot. Jap. 5: 155-169.
Inskipp, T. and Wells, S. 1979. International trade inwildlife. International Institute for Environment andDevelopment,London.
Instituto National para la Conservation de la Naturaleza(ICONA). 1992.Statusand conservationof the pardel
lynx (Lynx pardina) in the Iberian peninsula. Councilof Europe, Nature and Environment Ser. 55,
Strasbourg.
InternationalFur Trade Federation(IFTF). 1989.Key- factsReferences

aboutjurs. IFTF pamphlet, Holte, Denmark.
International Snow Leopard Trust (ISLT). 1993. Assessing
presence, relative abundance and habitat ofsnow leopard and their prey: a handbook offield techniques.
Seattle, Washington.
Ionescu, 0. 1993. Wildcat in Romania. Pp. 57-58 in Proc.
seminar on the biology and conservation of the wildcat (Felis silvestris), Nancy, France, 23-25 September
1992. Council of Europe, Strasbourg.
Iriarte, J.A., Johnson, W.E. and Franklin, W.L. 199 1.
Feeding ecology of the Patagonia puma in southernmost Chile. Revista Chilena de Historia Natural 64:
145-156.
-, Franklin, W.L., Johnson, W.E. and Redford, K.H.
1990. Biogeographic variation of food habits and bodysize of the American puma. Oecologia 85: 185-190.
Ishadov, N. 1983. Problems ofcaracal lynx ecology in
Turkmenia. In Redkie Vidy Mlekopitayuschikhi ikh
Okrana (Rare species of mammals and their conservation). Proc. of the 3rd All-Union Conf. Moscow, 1982,
pp. 105-107. Nauka Publ., Moscow, Russia.
IUCN-theWorld Conservation Union. 1976. Red data
book-terrestrial
mammals. IUCN, Gland, Switzerland
and Cambridge, U.K.
-. 1982. IUCN directory of neotropical protected areas.
Tycooly, Dublin.
*- . 1987a. IUCN directory of Afrotropical protectedareas. IUCN, Gland, Switzerland and Cambridge, U.K.
-. 1987b. IUCNpolicy
statement on captive breeding.
IUCN, Gland, Switzerland and Cambridge, U.K.
-. 1990a. IUCN directory of SouthAsianprotected areas.
IUCN, Gland, Switzerland and Cambridge,U.K.
- . 1990b. United Nations list of national parks and protected areas. IUCN, Gland, Switzerland andCambridge,U.K.
- . 1992a. Protected areas of the world: a review ofnational systems.Vol IV: Nearctic and Neotropical.
IUCN, Gland, Switzerland and Cambridge,U.K.
- . 1992b. Workshop abstracts *from the IVth World
Congress on National Parks and Protected Areas,
Caracas, Venezuela, I O-21 February 1992. IUCN,
Gland, Switzerland andCambridge,U.K.
-. 1994.1993 United Nations List of National Parks andProtected Areas. IUCN, Gland, Switzerland andCambridge,U.K.
- . 1995. IUCN Red List Categories. IUCN, Gland,
Switzerland.
- . In prep. Draft IUCN guidelinesfor the ecological sustainability of nonconsumptiveand consumptiveusesof
wild species.Paperpresentedat the 19th IUCN General
Assembly, BuenosAires, Argentina.
IUCN Environmental Law Centre. 1986.African wildlife
Zaws.IUCN EnvironmentalPolicy andLaw Occasional
Paperno. 3. IUCN, Gland, Switzerland andCambridge,

U.K.
IUCNAJNEPWWF. 1991. Caringfor the Earth: a strategy
for sustainableliving. IUCN-the World Conservation
Union, the United Nations Environmental Programme,
and WWF-the World Wide Fund for Nature. IUCN,
Gland, Switzerland andCambridge,U.K.
IUDZG/CBSG. 1993. The world zoo conservation strategy: the role of the zoos and aquaria of the world inglobal conservation. IUDZG-the
World Zoo
Organization and IUCN/SSC Captive BreedingSpecialistGroup. ChicagoZoological Society, Chicago.
Izawa, M., Doi, T. andY. Ono. 1989.Social systemof the
Iriomote cat (Felis iriomotensis).Abstractsfrom the 5th
intl. theriological congress 2: 608. Rome, 27-29
August.
-. 1991.Ecological study on the two speciesof Felidaein
Japan. In N. Maruyama et al., eds. WiZdZ$econservation: present trendsand perspectivesfor the 21st century. Proc. V Int. Congr. Ecol. Yokohama, Japan.
Jackson,P. 1987. Transfer of Siberian tiger from CITES
Appendix II to Appendix I. Cat News7: 15-16.BougyVillars, Switzerland.
- . 1989.A review by leopard specialistsof The statusof
the leopard in sub-SaharanAfrica by Martin and deMeulenaer. Information document no. 3 submitted tothe seventhmeetingof the Conferenceof the Partiesto
CITES, Lausanne,Switzerland.
-. 1991a. Endangered species: tigers. Apple Press,
London.
-. 1991b. Into the abyss- will the tiger follow the rhinos?
Cat News 15: 1. Bougy-Villars, Switzerland.
- . 1992a.The bonesof a dilemma. IUCN pressrelease,
Gland, Switzerland.
-. 1992b.International specialistsdiscussChina’sthreatened cats. Cat News 17: 9-10. Bougy-Villars,
Switzerland.
-. 1993a. The status of the tiger in 1993. Report to the
CITES Animals Committee,August 1993.Reproducedin Cat News 19: 5-l 1. Bougy-Villars, Switzerland.
-. 1993b.Tiger conservationmovesagainto centre stage.
Cat News 18: 2-3. Bougy-Villars, Switzerland.
Jackson,R. 1979. Snow leopardsin Nepal. ORYX
i 15:191195.
-. 1984. The snowleopard. Pp. 197-198in Theplight ofthe cats: Proc. of the meeting and workshop of the
IUCN/SSC Cat Specialist Group at Kanha National
Park, Madhya Pradesh, India, 9-12 April 1984.
Unpubl. report, IUCN/SSC Cat Specialist Group,
Bougy, Switzerland.
-. 1992. Snow leopard. Unpubl. data sheet,IUCN/SSC
Cat SpecialistGroup, Bougy-Villars, Switzerland.
- andAhlborn, G. 1984.A preliminary habitat suitability
model for the snow leopard (Panthera uncia) in West
Nepal.Intl. Ped. Book of SnowLeopards4: 43-52.

354

References- . 1988. Observations on the ecology of the snow leopard(Panthera uncia) in west Nepal. Pp.. 65-97 in H.
Freeman, ed. Proc. fifth international snow leopard
symposium. International Snow Leopard Trust andWildlife Institute of India, Conway, Bombay.
- . 1989. Snow leopards (Panthera uncia) in Nepal: homerange and movements. Nat. Geogr. Research 5(2): 161175.
- . 1990. The role of protected areas in Nepal in maintaining viable populations of snow leopards. Intl. Ped.
Book ofSnow Leopards 6: 5 l-69.
- 7Ahlborn, G., Ale, S., Gurung, D., Gurung, M. and U.
Yadav. 1994. Reducing livestock depredation in the
Nepalese Himalaya: case of the Annapurna Conservation Area. Unpubl. draft report to the U.S. Agency forInternational Development. Biosystems
Analysis,
Tiburon, California.
- and Hillard, D. 1986. Tracking the elusive snow leopard. Nat. Geog. 169(6): 793-809.
Jackson, R., Wang, Z.Y., Lu, X.D. and Y. Chen. 1994.
Snow leopards in the Qomolangma Nature Reserve of
the Tibet Autonomous Region. In J.L. Fox and Du
Jizeng, eds. Proc. of the seventh intl. snow leopard
symp. International Snow Leopard Trust, Seattle.
Jacobi, A. 1923. Zoologische Ergebnisse der Walter
Stotznerschen Expedition nach Szetschwan, Osttibet
und Tschili, auf Grund der Sammlungen andBeobachtungenDr. Hugo Weigolds.
2. Teil,
Mammalia. [Zoological findings of Walter Stotzner’s
expedition to Sichuan, eastern Tibet, and Chile, basedon the collections and observations of Dr Hugo
Weigold. Part 2, Mammalia.] Abh. Ber. MUX Tierkunde
Volkerkunde Dresden: 16.
Jalkotzy, M., Ross, I. and J.R. Gunson. 1992. Management
planfor cougars in Alberta. Wildl. Manage. Planning
Series 5, Alberta Forestry, Lands and Wildlife, Fish andWildl. Div., Edmonton.
James, W.E. In prep. Bobcat movements and habitat use
on Cumberland Island, Georgia during two years ofcontrolled releases. M.S. thesis, Univ. Georgia, Athens.
Janczewski, D.N., Yuhki, N., Gilbert, D.A., Jefferson,
G.T. and S.J. O’Brien. 1992. Molecular phylogenetic
inference from saber-toothed cat fossils of Ranch0 La
Brea. Proc. Natl. Acad. Sci. 89: 9769-9773.
Jardine, W. 1834. The naturalist’s library:w Mammalia,~012; the natural history of the Felinae. W.H. Lizars,
Edinburgh.
Jedrzejewski,
W., Schmidt, K., Milkowski,L.,
Jedrzejewska, B. and H. Okarma. 1993. Foraging bylynx and its role in ungulate mortality: the local(Bialowieza Forest) and Palearctic viewpoints. Acta
Theriologica 38(4): 385-403.
Jeffreys, A.J., Wilson, V. and S.L. Thein. 1985.
Hypervariable minisatellite regions in human DNA.

Nature 3 16: 76-79.
Jenny, D. 1993. Leopard research in Ivory Coast rain forest. Cat News 18: 12- 13. Bougy-Villars, Switzerland.
Jerdon, T.C. 1874. The mammals ofIndia: natural history.
John Wheldon, London.
Jewell, P.A. 1982. Conservation of the cheetah: should
cheetah be moved to distant areas? Unpubl. workshop
report, International Fund for Animal Welfare,
Cambridge.
Johns, A.D. 1986. Effects of habitat disturbance on therainforest wildlife in Brazilian Amazonin. Unpubl.
report, WWF, Washington, D.C.
- . 1989. Timber, the environment and wildlife in
Malaysian rain .forests. Final report to Institute ofSoutheast Asian Biology, Univ. Aberdeen, Aberdeen,
Scotland.
- . 1992. Species conservation in managed tropicalforests. Pp. 15-54 in T.C. Whitmore and J.A. Sayer,
eds. Tropical deforestation and species extinction.
Chapman and Hall, London.
Johnsingh, A.J.T. 1983. Large mammalian prey-predators
in Bandipur Tiger Reserve. J. Bombay Nat. Hist. Sot.
80: l-57.
- and Ravi Chellam. 1991. Asian lions. Pp. 92-93 in J.
Seidensticker and S. Lumpkin, eds. Great Cats.
Merehurst, London.
Johnsingh, A.J.T., Panwar, H.S. and Rodgers, W.A. 1991.
Ecology and conservation of large felids in India. Pp.
160-165 in N. Maruyama et al., eds. Wildlife conservation: present trends and perspectives for the 21st century. Proc. V Int. Congr. Ecol. Yokohama, Japan.
Johnson, K.A. 1989. Global CITES enforcement: any
progress? TRAFFIC USA Newsletter 9(4): 9- 11.
- . 1990. Bobcat (Felis rufus). Unpubl. data sheet,
IUCN/SSC/Cat
Specialist Group, Bougy-Villars,
Switzerland.
~ and T.K. Fuller. 1992. The leopard cat (Felis bengalensis) in China: ecology and management,for sustainable utilization. Unpubl. project proposal.
-, Yu, J.-P., Wang, Y.-X., Wang, S. and T.K. Fuller. 1993.
The leopard cat (Felis bengalensis) in China: ecologyI
and managementfor sustainable utilization. Unpubl.
progress report no. 1, CITES Secretariat, Geneva.
Johnson, K.G. and Pelton, M.R. 1981. A survey of procedures to determine relative abundance of furbearers inthe southeastern United States. Proc. Annu. Con5Southeast. Assoc. Fish Wildl. Agencies 35: 261-272.
Johnson, W.E. and Franklin, W.L. 199 1. Feeding and spatial ecology of Geoffroy’s cat (Felis geoflroyi) in southem Patagonia. J. Mamm. 72(4): 815-820.
- 7 and Iriarte, A. In press. Sympatric felid and canid
predators of Torres de1 Paine National Park, Chile. In
Patagonia Gem: the ecology and natural history - ofTorres de1 Paine National Pa-r-k, Chile.

355

References

Jones, M.L. 1977. Record keeping and longevity of felids
in captivity. Pp. 132- 138 in R.L. Eaton, ed. The world’s
cats 3: contributions to breeding, behavior, management and husbandry. Carnivore Res. Inst., Univ.
Washington, Seattle.
Jones, J.H. and Smith, N.S. 1979. Bobcat density and preyselection in central Arizona. J. Wildl. Manage. 43: 666672.
Jonsson, S. 1980. [Research and conservation of lynx inSweden.] Pp. 170-180 in A. Festetics, ed. [The Zynx inEurope.] Kilda, Greven (in German).
Jordan, D.B. 1990. Mercury contamination: another threat
to the Florida panther. End. Species Tech. Bull. 15(2): 1.
- . 1991. A proposal to establish a captive breeding population of Florida panthers. Final supplemental environmental assessment, Office of the Florida pantherrecovery coordinator, U.S. Fish and Wildlife Service,
Gainesville.
- . 1993. Participation schedule for the revised Florida
Panther Recovery Plan. U.S. Fish and Wildlife Service,
Gainesville, Florida.
- . 1994. Florida panther update: January-March 1994.
U.S. Fish and Wildlife Service, Gainesville, Florida.
Jorgenson, J.P. and Redford, K.H. 1993. Humans and big
cats as predators in the Neotropics. In N. Dunstone and
M.L. Gorman, eds. Mammals as predators. Proc.
Symp. 2001. Sot. Lund. 65. Clarendon, Oxford.
Joslin, P. 1973. The Asiatic lion: a study of ecology andbehaviour. Ph.D. thesis, University of Edinburgh,
Edinburgh.
- . 1984. The evironmental limitations and future of the
Asiatic lion. J. Bombay Nat. Hist. Sot. 8 1: 648-664.
- . 1988. A phototrapline for cold temperatures. Pp. 121182 in H. Freeman, ed. Proc. fifth international snow
leopard symposium. International Snow Leopard Trustand Wildlife Institute of India, Conway, Bombay.
- . 1990. Leopards in Iran. Pp. 13- 15 in A. Shoemaker, ed.
1989 Internatl.
leopard studbook.
Riverbanks
Zoological Park, Columbia, South Carolina.
Joubert, E., Morsbach, D. and Wallis, V. 1982. The 1982
distribution patterns and status of somemammalson
farms in South WestAfrica. Unpubl. report, Ministryof Wildlife Conservationand Tourism, Windhoek.
Kaczcensky, P. 1991. Untersuchungenzur Raumnutzung
weiblicher Luchse (Lynx lynx), sowie zur Abwanderung und Mortalittit ihrer Jungen im Schweizer Jura.
[Spatial organization of female lynx, and dispersalandmortality of their young, in the SwissJura Mountains.]
Dipl. thesis,Munich Univ., Munich.
Karami, M. 1992. Cheetah distribution in Khorasan
Province, Iran. Cat News 16: 4. Bougy-Villars,
Switzerland.
Karanth, K.U. 1987. Tigers in India: a critical review offield censuses.Pp. 118-132 in R.L. Tilson and U.S.

Seal,eds. Tigers of the world: the biology, biopolitics,
management and conservation of an endangered
species.Noyes, Park Ridge, New Jersey.
-. 1988. Analysis of predator-prey balance in Bandipur
Tiger Reserve with reference to census reports. J.
Bombay Nat. I-list. Sot. 85: l-8.
- . 1991. Ecology and managementof the tiger in tropical Asia. Pp. 156-159 in N. Maruyama et al., eds.
Wildlife conservation:present trendsand perspectives
for the 21st century. Proc. V Int. Congr. Ecol.
Yokohama, Japan.
- . 1993a.Predator-prey relationships among the large
mammalsof Nagarhole National Park (India). Ph.D.
thesis,Mangalore Univ., Mangalore.
- . 1993b. How many tigers? Field censusesin Inda.
Hornbill 1993 (1): 2-9. Bombay Nat. Hist. Sot.,
Bombay.
- and Sunquist,M.E. 1992.Population structure,densityand biomassof large herbivores in the tropical forestsof Nagarahole,India. J. Tropical Ecol. 8: 2 l-35.
Kartawinata, K. 1978.Biological changesafter logging inlowland dipterocarp forest. Biotrop. Special
Publication 3: 27-34.
Kashkarov, D.N. 1931. [Animals of Turkestan.] Tashkent(in Russian).
Keith, L.B. 1963. Wildlife’s ten year cycle. Univ. of
WisconsinPress,Madison.
-. 1974. Somefeaturesof population dynamics in mammals.Proc. 1Ith Internat. Congressof Game Biology,
Stockholm: 17-58.
-, Carey, J.R., Rongstad, O.J. and M.C. Brittingham.
1984. Demography and ecology of a declining snowshoehare population. Wildl. Monogr. 90: l-43.
Kesterson,M.B. 1988.Lynx homerange and spatial organization in relation to population density and prey
abundance.M.S. thesis,Univ. of Alaska, Fairbanks.
Khan, A.A. and Beg, M.A. 1986. Food of some mammalian predators in the cultivated areasof Punjab.
Pakistan J. 2002. 18(1): 7l-79.
Khan, J.A., Rodgers, W.A., Johnsingh, A.J.T. and P.K.
Mathur. 1990.Gir lion project: Ungulate habitat ecology in Gir. Unpubl. report, Wildlife Institute of India,
Dehra Dun.
Khan, Mohd. 1987. Tigers in Malaysia: prospectsfor the
future. Pp. 75-84 in R.L. Tilson and U.S. Seal, eds.
Tigers of the world: the biology, biopolitics, management and conservation of an endangered species.
Noyes, Park Ridge, New Jersey.
Khushwah,R.B.S. 1990.Thepast and presentproblem ofmanaging tigers with regard to past stock lifting andmankilling and later killing of tigers. Dipl. thesis,
Wildlife Institute of India, Dehra Dun.
Kier, A.B., Wightmann, S.R. and J.E. Wagner. 1982.
Interspeciestransmissionof Cytauxzoon felis. Am. J.

356

References

Vet. Rex 43: 101-105.
Kight, J. 1962. An ecological study of the bobcat Lynx
rufus (Schreber) in west-central South Carolina. M.S.
thesis, Univ. of Georgia, Athens.
Kiltie, R.A. 1984. Size ratios among sympatric neotropicalcats. Oecologia 61: 411-416.
Kimura, M. and Crow, J.F. 1963. The measurement ofeffective population number. Evolution 17: 279-288.
Kingdon, J. 1977. East African mammals: An atlas of evolution in Africa, vol. 3(A). Carnivores. Academic Press,
New York.
- . 1990. Arabian mammals: a natural history. Academic
Press, London.
Kinnear, N.B. 1920. The past and present distribution of
the lion in southeastern Asia. J. Bombay Nat. Hist. Sot.
27: 33-39.
Kitchener, AC. 199 1. The natural history of the wild cats.
Christopher Helm, London.
- . 1992. The Scottish wildcat Felis silvestris: decline andrecovery. Pp. 21-4 1 in Proc. Con& of theAssociationof
British Wild Animal Keepersand Ridgeway Trust for
EndangeredCats. Ridgeway Trust, Hastings,U.K.
- and Easterbee,N. 1992.The taxonomic statusof black
wild felids in Scotland. J. Zool., Lond. 227: 342-346.
Kitchener, A.C., McOrist, S. and J.M. Lynch. 1993.
Morphological and genetic discriminantsof European
wildcats, Felis silvestris, in Great Britain. Pp. 72-74 in
Proc. seminar on the biology and conservation of thewildcat (Felis silvestris), Nancy, France, 23-25
September1992. Council of Europe, Strasbourg.
Kitchings, J.T. and Story, J.D. 1984.Movement and dispersalof bobcatsin eastTennessee.J. Wildl. Manage.
48: 957-961.
Kleiman, D. 1974.The estrouscycle of the tiger. Pp. 60-75
in R.L. Eaton, ed. The world’s cats 2. World Wildlife
Safari, Winston, Oregon.
- and Eisenberg, J.F. 1973. Comparisonsof canid andfelid social systemsfrom an evolutionary perspective.
Anim. Behav. 21: 637-659.
Knick, ST. 1990.Ecology of bobcatsrelative to exploitation and a prey decline in southeasternIdaho. Wildl.
Monogr. 108: l-42.
- 7Sweeney, S.J., Alldredge, J.R. and J.D. B&tell. 1984.
Autumn and winter food habits of bobcats inWashington state.Great Basin Nat. 44: 70-74.
Kock, D. 1990.Historical record of a tiger, Panthera tigris(Linnaeus, 1758) in Iraq. Zoology in the Middle East
4: 11-15.
Koehler, G.M. 1990a.Population and habitat characteristics of lynx and snowshoe hares in north central
Washington.Can. J. 2001.68: 845-851.
- . 1990b.Canadalynx (Lynx canadensis).Unpubl. data
sheet,IUCN/SSC Cat SpecialistGroup, Bougy-Villars,
Switzerland.

-. 1991. Survey of the remaining population of South

China tigers. Unpubl. report, WWF International,
Gland, Switzerland.
- andB&tell, J.D. 1990.Managing spruce-fir habitat forlynx and snowshoehares.J. Forestry 88(10): 10-14.
Koehler, G.M. and Homocker, M.G. 1989. Influences ofseasons
on bobcatsin Idaho. J. Wildl. Manage. 53: 197202.
- . 1991. Seasonalresourceuse among mountain lions,bobcatsandcoyotes. J. Mamm. 72: 391-396.
Koford, C.B. 1973. Spotted cats in Latin America: aninterim report. Orq’x 12(l): 37-39.
- . 1976.Latin American Cats:economicvaluesandfuture
prospects.Pp. 79-88 in R.L. Eaton, ed. Theworld’s cats
3(l): contributions to status,managementand conservation. Carnivore Res.Inst., Univ. Washington,Seattle.
Konecny, M.J. 1989. Movement patternsand food habitsof four sympatric carnivore speciesin Belize, Central
America. Pp. 243-264 in K.H. Redford and J.F.
Eisenberg,eds.Advances in Neotropical mammalogy.
Sandhill Crane Press,Gainesville, Florida.
Korkishko, V.G. 1986. [Ecological features and behaviorof the Far Eastern leopard.] M.S. thesis, Univ.
Moscow, Moscow (in Russian).
- and Pikunov, D. 1994. Thepopulation number of the
Far East leopard in 1991 in Russia.Unpubl. report presentedto the SpeciesSurvival Commission,IUCN 19th
General Assembly, BuenosAires, Argentina.
Korneev, N.I. and Spasskaya,T. Kh. 1983. [The jungle
cat in Dagestan.] [Proc. Congr. All-Union Therio.
Sot.] 3: 64. Nauka, Moscow (in Russian).
Koshkarev, E.P. 1984. Characteristics of snow leopard(Uncia uncia) movementsin the Tien Shan. Intl. Ped.
Book of SnowLeopards4: 15-21.
- . 1989.[Snow leopardsin Kirgizia: statusand ecology.]
Nauka, Moscow (in Russian).
- . 1990. On the environment-related stability of snow
leopard (Uncia uncia) populationsin connection with
its location in the natural habitats and chances for
spread within the U.S.S.R. Intl. Ped. Book of Snow
Leopards6: 37-50.
Kosloff, L. and Trexler, T. 1987. The Convention on
International Trade in EndangeredSpecies:no carrot,
but where’sthe stick?Env. Law Reporter 17(7): 1022210236.
Koslowsky, J. 1904. DOSmamiferos de Patagonia cazadosen el valle de1Lago Blanc0 (Territorio de1Chubut).
[The mammalsof Patagoniahunted in the valley of theLago Blanc0 (Chubut Territory).] Revistade2Museo de
La Plata 11: 129-132 (in Spanish).
Kotwal, P.C. 1984. Lesserwild cats of Madhya Pradesh,
India. Pages95-99 in Theplight of the cats: Proc. of themeetingand workshopof the IUCNXSC Cat Specialist
Group at Kanha National Park, Madhya Pradesh,357

References(Lynx lynx) in Norway. Ph.D. thesis, Univ. Trondheim,
Trondheim.
- . 199 1. Reproduction in the European lynx, Lynx Lynx. 2.
Siiugetierk. 56: 146-158.
Labuschagne, W. 1979. [A bio-ecological and behaviouralstudy qf the cheetah, Acinonyx jubatus jubatus(Schreber, 1776).] M.S. thesis, Univ. Pretoria, Pretoria(in Afrikaans).
- . 1981. Aspects of cheetah ecology in the Kalahari
Gemsbok National Park. Paper presented at Internat.
Union of the Directorsof Zoological Gardens,
Washington, D.C.
Lacy, R.C. 1993. Vortex: a computer simulation model for
population viability analysis. Wildl. Res. 20: 45-65.
Laing, S.P. 1988. Cougar habitat selection and spatial
use patterns in southern Utah. M.S. thesis, Universityof Wyoming, Laramie.
Lamba, B.S. 1967. Occurrence of the desert cat, Felis
libyca ornata Gray near Poona. J. Bombay Nat. Hist.
Sot. 64(3): 552.
Lambert, M. 1966. A report on the Trinity College, Dublin
expedition to the High Atlas and Sahara. Trinity
College, Dublin.
Lande, R. 1979. Effective deme sizes during long-term
evolution estimated from rates of chromosomal
rearrangement. Evolution 33: 234-25 1.
- . 1988. Genetics and demography in biological conservation. Science 241: 1455- 1459.
- and Barrowclough,G.F. 1987. Effective population
size, genetic variation, and their use in population management. Pp. 87- 124 in M.E. Soul& ed. Viable populations jar conservation.
Cambridge Univ. Press,
Cambridge.
Lanly, J.P. 1982. Tropicaljorest resources. FAO Forestry
Paper 30, Rome.
Laurenson, M.K. 1992. Reproductive strategies in wild
female cheetahs. Ph.D. thesis, Univ. of Cambridge,
Cambridge.
-. 1993. Early maternal behavior of wild cheetahs: implications for captive husbandry. Zoo Biology 12: 3 l-43.
- . In press. Implications of high offspring mortality forcheetah population dynamics. In A.R.E. Sinclair and
P. Arease, eds. Serengeti II: research, conservation andmanagement ojan ecosystem. Univ. of Chicago Press,
Chicago.
- 7Caro, T. and M. Bomer. 1992. Female cheetah reproduction. Nat. Geog. Research 8(l): 64-75.
Lavauden, L. 1930. Essai sur l’histoire naturelle du lynx.
[Essay on the natural history of the lynx.] Imp. Allier,Grenoble.
Lawson, D. 1987. A survey ojthe eflects ojpredators on
sheepjarming in Natal. Unpubl. report, Univ. of Natal,
Pietermaritzburg.
Lay, D.M. 1967. A study of the mammals of Iran resulting

India, 9-12 April 1984. Unpubl. report, IUCN/SSC Cat
Specialist Group, Bougy, Switzerland.
Kowalski, K. and Rzebik-Kowalska, B. 1991. Mammals oj
Algeria. Polish Academy of Sciences, Warsaw.
Kratochvil, J. 1968. Survey of the distribution of populations of the genus Lynx in Europe. Acta. SC.nat. Brno 4:
5-12.
Kraus, D. and Marker-Kraus,
L. 199 1. The status of thecheetah (Acinonyx jubatus). Unpubl. data sheet,
IUCNISSC
Cat Specialist Group, Bougy-Villars,
Switzerland.
-. 1992. Cheetah Preservation Fund report. Cat News 17:
12- 14. Bougy-Villars, Switzerland.
Krebs, C.J. 1985. Ecology: the experimental analysis of
distribution and abundance, 3d edn. Harper and Row,
New York.
-, Boonstra, R., Boutin, S., Dale, M., Hannon, S., Martin,K., Sinclair, A.R.E., Smith, J.N.M. and R. Turkington.
1992. What drives the snowshoe hare cycle in Canada’s
Yukon? Pp. 886-896 in D.R. McCullough and R.H.
Barrett, eds. Wildlije 2001: populations. Elsevier, NewYork and London.
-, Gilbert, B.S., Boutin, S., Sinclair, A.R.E. and J.N.M.
Smith. 1986. Population biology of snowshoe hares. I.
Demography of food-supplemented populations in thesouthern Yukon, 1976- 1984. J. Anim. Ecol. 55: 963982.
Kruuk, H. and Turner, M. 1967. Comparative notes on predation by lion, leopard, cheetah and wild dog in the
Serengeti area, East Africa. Mammalia 3 1: l-27.
Kuhn, H.J. 1973. Zur Kenninis der Andenkatze, Felis(Oreailurus) jacobita Cornalia, 1865. [Knowledge of
the Andean cat.] Saugetierk. Mitt. 2 I(4): 359-364 (inGerman).
Kumerloeve, H. 1975. [The mammals (Mammalia) of
Syria and Lebanon.] Veriijj 2001. Staatssamml.
Munchen 18: 159-225 (in German).
Kunte, S. and Gore, A.P. 1986. On the probable error in a
census on waterholes. Hornbill 1986(3): 39-40.
Kurten, B. 1968. Pleistocene mammals of Europe.
Weidenfeld and Nicolson, London.
- . 1973. Geographic variation in size in the puma.
Comment. Biol. Sot. Sci. Fennica 62: l-8.
Kurten, B. and Rausch, R. 1959. Biometric comparisons
between North American and European mammals.
Acta. Arct. 11: 21-44.
- and Granqvist, E. 1987. Fossil pardel lynx (Lynx pardina speZaea Boule) from a cave in southern France.
Ann. 2001. Fennici 24: 39-43.
Kuryatkinov, A.L. 1983. [Numbers and the conservation
problems of rare mammals of northern Osetia.] [Proc.
Congr. All-Union Therio. Sot.] 3: 13. Nauka, Moscow(in Russian).
Kvam, T. 1990. Population biology of the European Lynx

358

Referencesfrom the Street expedition of 1962-63. Fieldiana 54.
- 7 Anderson, J.A.W. and J.D. Hassinger. 1970. New
records of small mammals from West Pakistan and
Iran. Mammalia 34: 100-106.
Le Berre, M. 1991. The role of Tassili N’ Ajjer (Algeria) in
the conservation of the great mammalian fauna in central Sahara. Pp. 18 I- 19 1 in J.A. McNeely and V.M.
Neronov, eds. Mammals in the palearctic desert: status and trends in the Sahara-Gobian region. RussianCommittee for the UNESCO programme on Man and
the Biosphere (MAB), Moscow.
Le Hien Hao. 1973. [Economic mammals of North
Vietnam]. Scientific and Technical House Press, Hanoi(in Vietnamese).
Le Houerou, H.N. and Gillet, H. 1986. Desertization in
African arid lands. Pp. 444-462 in M.E. Soul& ed.
Conservation biology: the science of scarcity anddiversity. Sinauer, Sunderland, Massachusetts.
Le Roux, P.G. and Skinner, J.D. 1989. A note on the ecology of the leopard (Panthera pardus Linnaeus) in the
Londolozi Game Reserve, South Africa. Afr. J. Ecol.
27: 167-171.
Leach, H.R. and Frazier, W.H. 1953. A study on the possible extent of predation on heavy concentrations ofvalley quail with special reference to the bobcat. Calzf
Fish and Game 39: 527-538.
Leal, R.P. 1979. Ensaios sobre a reproducae do jaguar em
cativeiro Panthera onca L. 1758 (Carnivora-Felidae).
[Notes on the reproduction of the jaguar in captivity.]
Acta. 2001. Lilloana 34: 107-l 12 (in Portugese).
Lee, A.R.
1992. Cheetah(Acinonyx
jubatus).
Management guidelines for the welfare ofzoo animals.
Federation of Zoological Gardens of Great Britain and
Ireland, London.
Lekagul, B. and McNeely, J.A. 1977. Mammals ofThailand. Association for the Conservation of Wildlife,
Bangkok.
Lembeck, M. 1978. Bobcat study, San Diego county,
Cal$ornia. Unpubl. report, California Dept. of Fish andGame, Sacramento.
- and Gould, G.I., Jr. 1979. Dynamics of harvested andunharvested bobcat populations in California. Bobcat
Res. Co@, Natl. Wildl. Fed. Sci. Tech. Ser. 6: 53-54.
Leopold, A.S. 1933. Game ‘management. Charles
Scribners Sons, New York.
- . 1959. Wildlife of Mexico. Univ. of Calif. Press,
Berkeley.
Leopold, B.D. and Krausman, P.R. 1986. Diets of three
predators in Big Bend National Park, Texas. J. Wildl.
Manage. 50: 290-295.
Leyhausen, P. 1963. [The South American spotted cats.] 2.
Tierpsychol. 20: 627-640 (in German).
- . 1977. Breeding endangered species. O~X 12(5): 427428.

-

the predatory and social behavior of domestic and wild cats. Garland, New York.
Engl. transl. B.A. Tonkin.
- and Falkena, M. 1966. Breeding the Brazilian ocelotcat Leopardus tigrinus in captivity. Int. Zoo Yearb. 6:
176-178.
Leyhausen, P. and Pfleiderer, M. 1994. The taxonomic
status of the Iriomote cat (Prionailurus iriomotensis[Imaizumi],1967). Cat News 2 1. Bougy-Villars,
Switzerland.
- and Tonkin, B. 1966. Breeding of the black-footed cat
in captivity. Int. Zoo Yb. 6: 178-182.
- and Wolff, R. 1959. Das Revier einer Hauskatze.
[Range of a housecat. ] 2. Tierpsychol. 16(6): 666-670.
Liao Yuanfa. 1988. [Some biological informations ofdesert cat in Qinghai.] Acta Theriologica Sinica 8(2):12% 13 1 (in Chinese with Engl. summary).
- and Tan Bangjie. 1988. A preliminary study on the geographical distribution of snow leopards in China. Pp.
5 l-63 in H. Freeman, ed. Proc. fifth international snow
leopard symposium.International Snow Leopard Trustand Wildlife Institute of India, Conway, Bombay.
Liberg, 0. and M. Sandell. 1988.Spatial organisationand
reproductive tacticsin the domesticcat andother felids.
Pp.. 83-98 in D.C. Turner and P. Bateson, eds. The
domestic cat: the biology of its behaviour. Univ.
CambridgePress,Cambridge.
Limoges, B. 1989.Resultsof the national wildlife inventory andproposalsfor modification qf the law on hunting. Unpubl. report, Ministry of Rural Development
andAgriculture, Republic of Guinea-BissauandIUCN.
Lin, H. and Chen, S. 1988. [Original colour atlasfor discriminating Chinesetraditional drugs.] Science andTechnology Press, Guangdong (in Chinese withEnglish introduction).
Lindburg, D.G., Durrant, B.S., Millard, S.E. and J.E.
Oosterhuis.1993.Fertility assessmentof cheetahmaleswith poor quality semen.Zoo Biology 12 (1): 97- 104.
Lindemann, W. 1953. Einiges Uber die Wildkatze derOstkarpaten(Felis s. silvestris Schreber,1777). [Someremarkson wildcats in the East Carpathianmountains.]
Saugetierk.Mitt. 1: 73-74.
Linden, E. 1994. Tigers on the brink. Time magazine 28
March: 50-57.
Lindeque, P., Preisser,T., Nowell, K., Brain, C., and P.
Turnbull. In prep. Anthrax in wild cheetahs in the
EtoshaNational Park.
Lindzey, F.G. 1987. Mountain lion. Pp. 656-668 in M.
Novak, J. Baker, M. Obbard, andB. Malloch, eds.Wild
furbearer management and conservation in North
America. Ontario Ministry of Natural Resources,
Toronto, Canada.
-, Van Sickle, W.D., Laing, S.P.andC.S. Mecham. 1992.
Cougar population responseto manipulation in south-359. 1979. Cat behavior:References

447-461.
Lukarevsky, V.S. 1990. [The presentstatusof the leopard
population in Turkmenia.] [Proc. Congr. AU- Union
Therio. Sot.] 5: 160-161.Nauka, Moscow (in Russian).
- . 1993. [Leopard, Panthera pardus, in the western
Kopetdag: distribution, numbers,ecology and behavior.] 2001.Zhurn. 72(l) (in Russian).
Lydekker, R. 1896.A handbook to the Carnivora. LloydLtd., London.
Lynch, C.D. 1983.The mammalsof the OrangeFree State.
Mem. nas.Mus., Bloemfontein 18: I-2 18.
Ma Yiqing. 1986. [The Pantherines.] Pp. 198-200 in[Mammals of Heilongjiang province.] Heilongjiang
Scienceand Technology Press,Harbin (in Chinesewith
English summary).
Maberly, C.T.A. 1966. Animals of East Africa, 2d edn.
Hawkins, Nairobi.
MacClintock, D. 1966. Chinchilla. Pacific Discovery”
19(3): 18-25.
MacDonald, D.W., Kerby, G. and W.C. Passanisi.1991.
Wild catsandferal cats.Pp. 162-169in J. Seidensticker
and S. Lumpkin, eds.Great cats. Merehurst, London.
Mace, G.M. 1993.The statusof proposalsto redefine the
IUCN Threatened SpeciesCategories.Pp. xlviii-lv in
B. Groombridge, ed. 1993. 1994 IUCN Red List of
threatened animals. IUCN, Gland, Switzerland andCambridge,U.K.
- and Lande, R. 1991. Assessing extinction threats:
toward a reevaluation of IUCN threatenedspeciescategories.Conserv.Biol. 5(2): 148-157.
Mace, G. and Stuart, S. 1994. Draft IUCN Red List
Categories, Vers. 2.2. Species21-22: 13-24. IUCN,
Gland, Switzerland.
Macey, A. 1979.Easterncougar, Felis concolor. Pamphletby the Committeeon the Statusof EndangeredWildlifein Canada,Government of Canada.
MacKinnon, J. 1990. Review of the nature conservation
system,national parks and protected areas. Vietnam
Tropical Forestry Action Plan, Forestry SectorReview.
Unpubl. report, United Nations Food and Agriculture
Organization, Hanoi.
- and MacKinnon, K. 1986a. Review of the protectedareas systemof the Afrotropical realm. IUCN, Gland,
Switzerland and Cambridge,U.K.
- . 1986b. Review of the protected areas svstemof theIndomalayan realm. IUCN, Gland, Switzerland andCambridge,U.K.
- 7Child, G. and J. Thorsell. 1986. Managing protectedareas in the tropics. IUCN, Gland, Switzerland andCambridge,U.K.
MacPherson, N. and Fernando,B.K. 1991. Opportunities
for improved environmental management in
Afghanistan. Draft unpubl. report, Office of the Coordinator for United Nations Humanitarian andern Utah. Wildl. Sot. Bull. 20: 224-227.
Linnaeus, C. 1758. Systema Naturae,
10th edn.
Stockholm.
Litvaitis, J.A., Clark, A.G. and J.H. Hunt. 1986. Prey selection and fat deposits of bobcats (Felis rujfus) during
autumn and winter in Maine. J. Mamm. 67: 389-392.
-, Major, J.T. and J.A. Sherburne. 1987. Influence of season and human-induced mortality on spatial organization of bobcats (Felis rufus) in Maine. J. Mamm. 68( 1):
IOO- 106.
Litvaitis, J.A., Stevens, C.L and W.W. Mautz. 1984. Age,sex and weight of bobcats in relation to winter diets. J.
Wildl. Manage. 48: 632-635.
Locke, A. 1954. The tigers of Trengganu. Museum Press,
London.
Logan, K.A. and Irwin, L.L. 1985. Mountain lion habitats
in the Big Horn Mountains, Wyoming. WiZdZ.Sot. Bull.
13: 257-262.
-, and R. Skinner. 1986. Characteristics
of a hunted
mountain lion population in Wyoming. J. Wildl.
Manage. 50(4): 648-654.
Logan, T., Eller, A.C., Morrell, R., Ruffner, D. and J.
Sewell. 1993. Florida panther habitat protection plan.
Unpubl.
report,FloridaPanther Interagency
Committee, U.S. Fish and Wildlife Service.
Loudon, A.S.I. 1985. Lactation and neonatal survival inmammals. Symp. Zool. Sot. Lund. 54: 183-207.
Low, J. 199 1. The smuggling of endangered wildlife
across the Taiwan Strait. TRAFFIC International,
Cambridge.
Lowery, G.H., Jr. 1936. A preliminary report on the distribution of the mammals of Louisiana. Proc. Acad. Sci.
3: 1l-39.
Lozan, M.N. and Korcmar, N.D. 1965. [Materials on theecology of predators in the riverine thickets of the Prutriver.] Ochrana prirody MoZdavii 3: 140- 149 (inRussian).
Lu Houji. 1987. Habitat availability and prospects fortigers in China. Pp. 7 l-74 in R.L. Tilson and U.S. Seal,
eds. Tigers qf the world: the biology, biopolitics, management and conservation of an endangered species.
Noyes, Park Ridge, New Jersey.
- and Sheng Helin. 1986a. Distribution and status of theChinese tiger. Pp. 5 l-58 in S.D. Miller and D.D.
Everett, eds. Cats of the world: biology, conservation,and management. National Wildlife Federation,
Washington,D.C.
- and Sheng Helin. 1986b. The status and population
fluctuation of the leopard cat in China. Pp. 59-62 in
S.D. Miller and D.D. Everett, eds. Cats of the world:
biology, conservation, and management. National
Wildlife Federation,Washington, D.C.
Ludlow, M.E. and Sunquist, M.E. 1987. Ecology andbehavior of ocelotsin Venezuela.Nat. Geog.Res.3(4):360References

Economic
Assistance
Programmes
relating to
Afghanistan, Geneva.
Maehr, D.S. 1990. The Florida panther and private lands.
Conserv. Biol. 4(2): 167- 170.
- and Brady, J.R. 1986. Food habits of bobcats in Florida.
J. Mamm. 67(l): 133-138.
-, Land, E.D. and M.E. Roelke. 199 1a. Mortality patternsof panthers in southwest Florida. Proc. Ann. Conf
Southeast Fish Wildl. Agencies 45.
- 7Land, E.D., and J.C. Roof. 1991b. Social ecology ofFlorida panthers. Nat. Geog. Res. 7: 4 14-43 1.
Makacha, S. and Schaller, G.B. 1969. Observations onlions in the Lake Manyara National Park, Tanzania. E.
Afr. Wildl. J. 7: 99- 103.
Makombe, K., ed. 1994. Sharing the land: wildlife, people and development in Africa. IUCN/ROSA Environmental Issues Series No. 1, IUCN/ROSA,
Harare,Zimbabwe and IUCN/SUWP, Washington, D.C.
Malafeev, Y.M. and Kryazhimskiy, F.V. 1984. [European
lynx Lynx lynx feeding and relationships with ungulates
in the central Urals, U.S.S.R..] Bjull. Mosk. obsc. ispty.
prir., otd. biol. 89: 70-81 (in Russian).
Mallon, D. 1984a. The snow leopard in Ladakh. Intl. Ped.
Book ofSnow Leopards 4: 23-37.
- . 1984b. The snow leopard, Panthera uncia, in
Mongolia. Intl. Ped. Book of Snow Leopards 4: 3- 10.
- . 1985. The mammals of the Mongolian Peoples
Republic. Mammal Review 15: 71-102.
- . 1987. An ecological survey of the snow leopard in
Ladakh. MS. thesis, Univ. of Manchester, Manchester.
- . 199 1. Status and conservation of large mammals in
Ladakh. Biol. Conserv. 56: 101-119.
Mallory, F.E. 1987. Impact of prey on furbearer predators. Pp. 141- 150 in M. Nowak, J. Baker, M. Obbard
and B. Malloch, eds. Wildfurbearer management andconservation in North America. Ministry of Natural
Resources, Toronto, Ontario.
Mambetzhumaev,
A.M. and Paliangazov, M. 1968.
[Ecology, distribution and practical value of somespecies of cats (Carnivora, Felidae) in the Karakalpak
ASSR.] 2001. Zh. 47: 423-431 (in Russian).
Manzani, P.R. and Monteiro F., E.L.A. 1989. Notes on
the food habits of the jaguarundi, Felis yagouaroundi(Mammalia: Camivora). Mammalia 53: 659-660.
Mares, M.A. and Ojeda, R.A. 1984. Fauna1 commercialization and conservation in South America. Bioscience
34(9): 580-584.
- and M.P. Kosco. 198 1. Observations on the distribution
and ecology of the mammals of Salta Province,
Argentina. Ann. Carnegie Mus. 50: 15 l-206.
Marker, L. and S. O’Brien. 1989. Captive breeding of thecheetah (Acinonyxjubatus)
in North American zoos
(1871-1986). Zoo Biology 8: 3-16.
Marker-Kraus,
L. 1992. International cheetah studbook

1991. NOAHS Centre, National Zoological Park,
Washington, D.C.
- 7Farrington, M., Kraus, D., Henkel, U. and D. Bounds.
1990. International cheetah studbook questionnairesummary. NOAHS Center, National Zoological Park,
Washington, D.C.
- and Grisham, J. 1993. Captive breeding of cheetahs inNorth American zoos. Zoo Biology 12(l): 5-18.
- and Kraus, D. 1991. 1991 Annual Report. Unpubl.
report, Cheetah Conservation Fund, Windhoek.
Marston, M.A. 1942. Winter relations of bobcats to whitetailed deer in Maine. J. Wildl. Manage. 6: 328-337.
Martin, E.B. 1987. Deadly love potions. Animal Kingdom90(l): 16-21.
- 1992a. The trade and uses of wildlife products in Laos.
‘TRAFFIC Bulletin 13( 1): 23-28.
- . 1992b. Observations on the wildlife trade in Viet Nam.
TRAFFIC Bulletin 13(2): 61-67.
- . 1992~. The poisoning of rhinos and tigers in Nepal.
Oryx 26: 82-86.
Martin, R.B. 1986. Communal Areas Management
Programme for Indigenous Resources (CAMPFIRE).
Government Printers, Harare.
-and de Meulenaer, T. 1988. Survey of the status of theleopard (Panthera pardus) in sub-Saharan Africa.
CITES Secretariat, Lausanne.
Maruska, E., Roychoudhury, A.K., Simmons, L.G. and K.
Latinen. 1987. White tiger politics. Pp. 372-395 in in
R.L. Tilson and U.S. Seal, eds. Tigers of the world: thebiology, biopolitics, managementand conservation of.
an endangeredspecies.Noyes, Park Ridge,New Jersey.
Masua, R., Michihiro, C., Shinyashiki, F. and Bando, G.
1994.Molecular phylogenetic statusof the Iriomote Cat
Felis iriomotensis inferred from mitochondrial DNA
sequenceanalysis.Zoological Science11:597-604.
Mather, A.S. 1990. Global forest resources.BellhavenPress,London.
Matjuschkin, E.N. 1978. [The lynx.] A. Ziemsen,
Wittenberg (in German).
-, Zhivotchenko, V.I. andE.N. Smimov. 1980.TheAmur
tiger in the U.S.S.R. Unpubl. report, IUCN, Gland,
Switzerland.
Matson, J.R. 1948.Cat kills deer. J. Mamm. 29: 69-70.
May, D.W. 1981. IIabitat utilization by bobcats in eastern Maine. M.S. thesis,Univ. Maine, Orono.
May, R.M. 1988.Conservationanddisease.Consent.Biol.
2( 1): 28-29.
- . 1990.Taxonomy asdestiny. Nature 347: 129-130.
Mayr, E. 1940. Speciation phenomenain birds. Am. Nat.
74: 249.
- . 1963.Animal speciesand evolution. Harvard Univ.
Press,Cambridge.
- . 1970. Populations, speciesand evolution. Belknap
Press,Cambridge.

361

References

Merehurst, London.
1994.Lessonsfrom the past: forests and biodiversity.
Biodiversity and Conservation3: 3-20.
- and Miller, K.R. 1984. National parks, conservation,and development: the role of protected areas in sustaining society. Proceedingsof the World Congresson
National Parks, Bali, Indonesia, 11-22 October 1982.
SmithsonianInstitution Press,Washington,D.C.
McOrist, S., Boid, R., Jones, T.W., Easterbee, N.,
Hubbard,A.L. and0. Jarrett. 1991. Someviral andprotozool diseasesin the European wildcat. J. Wildl.
Diseases27: 693-696.
McVittie, R. 1979. Changes in the social behaviour ofSouth West African cheetah.Madoqua 2(3): 171-184.
Mech, L. 1980. Age, sex, reproduction, and spatial organization of lynxes colonizing northeasternMinnesota.
J. Mamm. 61: 26l-267.
Medway, G. 1965.Mammalsof Borneo.$eld keysand anannotated checklist. Royal Asiatic Society, Kuala
Lumpur.
-. 1978. The wild mammals of Malaya (Peninsular
Malaysia) and offshoreislandsincluding Singapore,2dedn. Oxford Univ. Press,Kuala Lumpur.
Mehrer, C.F. 1975.Someaspectsof reproduction in captive mountain lions Felis concolor, bobcat Lynx rufus,
and lynx Lynx canadensis.Ph.D. thesis, Univ. North
Dakota, Grand Forks.
Mehta, J.N. and Dhewaju,R.G. 1990.A note on the recordof cloudedleopardsin Nepal. Tiger Paper 17(l): 21-22.
Melisch, R. 1995.Early record offishing cat in Peninsular
Malaysia. Cat News 22. Cat SpecialistGroup, Bougy,
Switzerland.
Melisch, R., Asmoro, P., Lubis, I. and Kusumawardhani,
L. 1995. Distribution and status of the fishing cat,Prionailurus viverrinus rhizophoreus, in West Java,
Indonesia.Faun. Abh. Tierkd. Mus. 20. Dresden.
Mellen, J. 1989. Reproductive behavior of small captive
cats (Felis ssp.).Ph.D. thesis,Univ. Calif., Davis.
-. 1991. Factors influencing reproductive successin
small captive exotic felids (Felis spp.j: a multiple
regressionanalysis.Zoo Biol. 10: 95-l 10.
-. 1992. Effects of early rearing experience on subsequent adult sexualbehaviour usingdomesticcats (Felis
catus) asa model for exotic smallfelids. Zoo Biol. 11:
17-32.
-. 1993.A comparative analysisof scent-marking,socialand reproductive behavior in 20 speciesof small cats
(Felis). American Zoologist 33.
-, Cheney, C. and J. Barker. 1992. A summaryof camivore spacein AAZPA zoos: 1992.Pp. 117-148in D.E.
Wildt, J.D. Mellen and U.S. Seal, eds. 1992. Felid
action plan, I991 and 1992: AAZPA Felid Taxon
Advisory Group regional collection plan and IUCN
Captive Breeding SpecialistGroup globalfelid captive

Mazak, V. 1979. [The tiger, 1st edn.] Ziemsen,
Wittenberg/Lutherstadt (in German).
-. 1981. Panthera tigris. Mammal. Species 152: 1-8.
McBride, C.J. 1977. The white lions of Timbavati.
Paddington Press, London.
- . 1984. Age and size categories of lion prey in Chobe
National Park, Botswana. Botswana Notes and Records
16: 139-143.
-. 1990. Liontide. Jonathan Ball, Johannesburg.
McBride, R.T. 1976. The status and ecology of the mountain lion Felis concolor stanleyana ofthe Texas-Mexico
border. M.S. thesis, Sul Ross State Univ., Alpine,
Texas.
McCarthy, T.J. 1992. Notes concerning the jaguarundi cat(Herpailurus yagouaroundi) in the Caribbean lowlandsof Belize and Guatemala. Mammaha 56: 302-306.
McCloskey, J.M. and Spalding, H. 1989. A reconnaissance-level inventory of the amount of wilderness
remaining in the world. Ambio 18(4): 221-227.
McCord, C.M. 1974. Selection of winter habitat by bobcats on the Quabbin Reservation, Massachusetts. J.
Mamm. 55: 428-437.
- and Cardoza, J.E. 1982. Bobcat and lynx. Pp. 728-766
in J.A. Chapman and G.A. Feldhamer, eds. Wild mammals ofNorth America: biology, management and economics. John Hopkins Univ. Press, Baltimore.
McDougal, C. 1977. The face of the tiger. Rivington
Books and Andre Deutsch, London.
-. 1987. The man-eating tiger in geographical and historical perspective. Pp. 435-448 in R.L. Tilson and U.S.
Seal, eds. Tigers ofthe world: the biology, biopolitics,
management, and conservation of an endangered
species. Noyes, Park Ridge, New Jersey.
-. 1991. Chuchchi: the life of a tigress. P 104 in J.
Seidensticker
and S. Lumpkin, eds. Great cats.
Merehurst, London.
McKeown, S. 1992. Joint management of species cheetah
breeding programme. Pp. 78-88 in P. Mansard, ed.
Cats: proc. conference/workshop
held at Chester Zoo
on October 10, 1992 by the Ridgeway Trust for
Endangered Cats and the Association of British Wild
Animal Keepers.Ridgeway Trust for EndangeredCats,
Hastings,Sussex,U.K.
McLaughlin, R. 1970.Aspects of the biology of cheetahs
Acinonyx jubatus(Schreber)in Nairobi National Park.
M.S. thesis,Univ. of Nairobi, Nairobi.
McMahan, L. 1986.The internationalcat trade.Pages461488 in S.D. Miller and D. Everett, eds. Cats of theworld: biology, conservation and management.
National Wildlife Federation,Washington, D.C.
McNab, B.K. 1971. On the ecological significance of
Bergmann’srule. Ecol. 52: 845-854.
McNeely, J.A. 1991.Do wild cats have a future? Pp. 222231 in J. SeidenstickerandS. Lumpkin, eds.Great cats.

-.

362

References

action plan. National Zoological Park, Front Royal,
Virginia.
Melquist, WE. 1984. Status survey ofotters (Lutrinae)
and spotted cats (Felidae) in Latin America. Unpubl.
report, Univ. of Idaho, Moscow and IUCN, Gland.
Mendelssohn, H. 1989. Felids in Israel. Cat News 10: 2-4.
Bougy-Villars, Switzerland.
Menotti-Raymond, M. and O’Brien, S.J. 1993. Dating thegenetic bottleneck of the African cheetah. Proc. Natl.
Acad. Sci. 90.
Miller, A.M., Roelke, M.E., Goodrowe, K.L., Howard,
J.G. and D.E. Wildt. 1990. Oocyte recovery, maturationand fertilization in vitro in the puma (Felis concolor). J.
Reprod. and Fertility 88: 249-258.
Miller, D.J. and Jackson, R. 1994. Livestock and snow
leopards: making room for competing users on the
Tibetan Plateau. In J.L. Fox and Du Jizeng, eds. Proc.
ofthe seventh intl. snow leopard symp. International
Snow Leopard Trust, Seattle.
Miller, S.D. 1980. The ecology of the bobcat in south
Alabama. Ph.D. thesis, Auburn University, Auburn,
Alabama.
- and Rottman, J. 1976. [Guide to the recognition ofChilean mammals.] Editora National Gabriela Mistral,
Santiago (in Spanish).
- and Speake, D.W. 1978. Prey utilization by bobcats onquail plantations in southern Alabama. Proc. Annu.
Conf Southeast Assoc. Fish and Wildl. .4gencies 32:
100-l 11.
Milliken, T., Nowell, IS. and J. Thomsen. 1993. The
decline of the black rhino in Zimbabwe: implications
for-future rhino conservation. TRAFFIC International,
Cambridge, U.K.
Millington, S.J. and T. Anada. 199 1. Biological diversity
assessment for Niger. Unpubl. report, WWF-Niger,
Niamey.
Mills, J.A. 1993. Report to the CITE’S Secretariat/Standing
Committee: a brief look at the tiger bone trade in South
Korea. TRAFFIC International, Cambridge.
Mills, M.G.L. 1984. Prey selection and feeding habits of
the large carnivores in the southern Kalahari. Koedoe
Suppl. : 28 l-294.
- . 1990. Kalahari hyaenas: the comparative behavioural
ecology of two species. Unwin-Hyman, London.
- . 199 1. Conservation management of large carnivores inAfrica. Koedoe 34( 1): 8 l-90.
- and Biggs, H.C. 1993. Prey apportionment and related
ecological relationships between large carnivores in
Kruger National Park. In N. Dunstone and M.L.
Gorman, eds. Mammals as predators. Proc. Symp.
2001. Sot. Lond. 65. Clarendon, Oxford.
-, Nel, J.A.J. and J.du P. Bothma. 1984. Notes on somesmaller carnivores from the Kalahari Gemsbok
National Park. Koedoe Suppl. : 22 l-227.

- and Shenk, T.M. 1992. Predator-prey relationships: theimpact of lion predation on wildebeest and zebra populations. J. Anim. Ecol. 61: 693-702.
- 7 Wolff, P., Le Riche, E.A.N. and I.J. Meyer. 1978.
Some population characteristics of the lion Panthera
Zeoin the Kalahari Gemsbok National Park. Koedoe 2 1:
163-171.
Misonne, X. 1959. [Zoogeographic analysis of the mammals of Iran.] Memoires d’btstitut Royal des Sciences
Naturelles de Belgique 59(2) (in French).
Mitchell, B., Shenton, J. and J. Uys. 1965. Predation onlarge mammals in the Kafue National Park, Zambia.
2001. Africana 1: 297-3 18.
Miththapala, S. 1992. Genetic and morphological variation in the leopard (Panthera pardus): a geographicallywidespread
species. Ph.D. thesis, Univ. Florida,
Gainesville.
- 7Seidensticker, J., Phillips, L.G., Fernando, S.B.U. and
J.A. Smallwood. 1989. Identification of individual
leopards (Panthera pardus kotiya) using spot patternvariation. J. Zool., Lond. 218: 527-536.
Mizutani, F. 1993. Home range of leopards and theirimpact on livestock on Kenyan ranches. In N. Dunstone
and M.L. Gorman, eds. Mammals as predators. Proc.
Symp. 2001. Sot. Lond. 65. Clarendon, Oxford.
Mohapatra, S. 1988. No let-up in killing of wildlife.
Hindustan Times (New Delhi), 21 April.
Mondolfi, E. 1986. Notes on the biology and status of thesmall wild cats in Venezuela. Pp. 125- 146 in S.D.
Miller and D.D. Everett, eds. Cats of the world: biology, conservation and management. National Wildlife
Federation, Washington, D.C.
- and Hoogesteijn, R. 1986. Notes on the biology and status of the jaguar in Venezuela. Pp. 85-124 in S.D.
Miller and D.D. Everett, eds. Cats of the world: biology, conservation and management. National Wildlife
Federation, Washington, D.C.
Monod, T. 1965. Comment-Discussion
Section. Pp. 547654 in Howell, F.C. and Bourliere, F. (eds.). African
ecology and human evolution. Methuen, London.
Moolman, L.C. 1984a. There’s a trick to catching a caracal. Custos 13(8): 42-43.
- . 1984b. [Comparison of feeding habits of the caracal
Felis caracal inside and outside of the Mountain Zebra
National Park.] Koedoe 27: 12 1- 129 (in Afrikaans).
- . 1986. [Aspects of the ecology and behavior of the caracal Felis caracal Schreber, I776 in the Mountain Zebra
National Park and surrounding farms.] M.S. thesis,
Univ. Pretoria, Pretoria (in Afrikaans).
Moore, D. 1992. Re-establishinglarge predators.
Reintroduction News 4: 6.
Morrison-Scott,
T.C.S. 1952. The mummified cats ofancient Egypt. Proc. 2001. Sot. Lond. 121: 86 l-867.
Morsbach, D. 1984-1986. [The behavioral ecology and363

References

Alaska lynx. M.S. thesis, Univ. Alaska, Fairbanks.
Navarro, D.L. 1985. Status and distribution of the ocelot(Felis pardalis) in south Texas. M.S. thesis, Texas A &I Univ., Kingsville.
Nawa, A. 1968. [Observations on the breeding habits andthe growth of Prionailurus bengalensis manchurica in
captivity.] J. Mamm. Sot. Japan 4: 12-19 (in Japanesewith English summary).
Negi, S.S. 1969. Transplanting of Indian lion in Uttar
Pradesh state. Cheetal 12: 98-101.
Nellis, C.H., Wetmore, S.P. and L.B. Keith. 1972. Lynxprey interactions in central Alberta. J. Wildl. Manage.
36: 320-329.
Neronov, V.M. and Bobrov, V.V. 1991. Conservation ofrare mammals in deserts of the U.S.S.R. Pp. 232-241
in J.A. McNeely and V.M. Neronov, eds. Mammals in
the Palearctic desert: status and trends in the SaharaGobian region. Russian Committee for the UNESCO
programme on Man and the Biosphere (MAB),
Moscow.
Nevo, E. 1978. Genetic variation in natural populations:patterns and theory. Theoretical Population Biology 13:
121-177.
Newman, A., Bush, M., Wildt, D.E., van Dam, D.,
Frankehuis, M., Simmons, L., Phillips, L. and S.J.
O’Brien. 1985. Biochemical genetic variation in eight
endangered or threatened feline species. J. Mamm.
66(2): 256-267.
Nguyen Xuan Dang and Pham Trong Ath. 1992. Tigers
threatened in Vietnam. Cat News 16:5. Bougy-Villars,
Switzerland.
Nichols, D.G., Fuller, KS., McShane-Caluzi, E., and E.
Klerner-Eckenrode.
199 1. Wildlife trade laws of Asiaand Oceania. A.L. Gaski and G. Hemley, eds. TRAFFIC USA, WWF-U.S., Washington, DC.
Nilsson, G. 1980. Facts about fur. Animal Welfare
Institute, Washington,D.C.
Noiseux, F. andDoucet, G.J. 1987. [Study of the Canadianlynx population in the Laurentides Fauna1 Reserve,
Quebec.] Unpubl. report, Quebec Minis&e du Loisir,de la Chasse,et de la P&he, Montreal (in French).
Norton, P.M. 1984.Leopardconservationin SouthAfrica.
Afr. Wildlife 38: 191-196.
-. 1986.Historical changesin the distribution of leopardsin the CapeProvince, South Africa. Bontebok 5: l-9.
- . 1990.How many leopards?A criticism of Martin andde Meulenaer’spopulation estimatesfor Africa. S.Afr.
J. Sci. 86: 218-220.
- and Henley, S.R. 1987. Home range and movementsof male leopards in the Cedarberg Wilderness Area,
CapeProvince. S.Afr. J. Wildl. Res. 17: 4 l-48.
- andA.B. Lawson. 1985.Radiotracking of leopardsand
caracalsin the Stellenboscharea,CapeProvince. S.Afr.
J. Wildl. Res. 15(l): 17-24.

movement of cheetahs on farmland in Southwest
Africa/Namibia.] Annual progress reports submitted to
the Directorate of Nature Conservation and Recreation
Resorts, Government of Namibia, Windhoek (inAfrikaans).
-. 1987. Cheetah in Namibia. Cat News 6: 25-26. BougyVillars, Switzerland.
Mowat, G. 1993. Lynx recruitment during a snowshoe
hare population peak and crash. M.S. thesis, Univ. ofAlberta, Edmonton.
Muckenhim, N.A. and Eisenberg, J.F. 1973. Home ranges
and predation of Ceylon leopard. Pp. 142- 175 in R.L.
Eaton, ed. The world’s cats l(1). World Wildlife Safari,
Winston, Oregon.
Mulliken, T. and Thomsen, J. 1990. U.S. bird trade: thecontroversy continues although imports decline. TRA FFIG USA Newsletter 1O(3): 1- Il.
Munson, L. and Mason, R. 199 1. Pathological findings in
the uteri of progestagen-implanted exotic felids. P 3 11 inProc. Amer. Assoc. Zoo Vet. Congr. Calgary, Alberta.
Muntyanu, A.I., Vasiliev, A.G. and P.T. Chegorka. 1993.
Distribution and number of the European wildcat (Felis
silvestris Schreber, 1777) in R. Moldova. Pp. 59-61 in
Proc. seminar on the biology and conservation of thewildcat (Felis silvestris),
Nancy, France, 23-25
September 1992. Council of Europe, Strasbourg.
Murphy, KM. 1983. Relationships between a mountainlion population and hunting pressure in western
Montana. M.S. thesis, Univ. of Montana, Missoula.
Muul, I. and Lim, B.L. 1970. Ecological and morphological observations of Felis planiceps. J. Mamm. 5 l(4):806-808.
Myers, N. 1973. The spotted cats and the fur trade. Pp.
276-323 in R.L. Eaton, ed. The world’s cats 1. World
Wildlife Safari, Winston, Oregon.
- . 1975. The cheetah Acinonyx jubatus in Africa. IUCN
Monograph No. 4. Morges, Switzerland.
- . 1976. The leopard Panthera pardus in Africa. IUCN
Monograph No. 5. Morges, Switzerland.
- . 1986. The small cats of Africa. Pp. 192- 199 in Birds
and mammals.
National
Wildlife
Federation,
Washington, D.C.
- . 1989. Deforestation rates in tropical forests and theirclimatic implications. Unpubl. report, Friends of the
Earth, London.
Nader, I.A. 1989. Rare and endangered mammals of Saudi
Arabia. Pp. 220-233 in A.H. Abu-Zinada, P.D. Goriup
and LA. Nader, eds. Wildlife conservation and development in Saudi Arabia. National Commission for
Wildlife Conservation and Development Publ. No. 3,
Riyadh.
Nasilov, S.B. 1972. [Diet of the wildcat in Azerbaijan.]
2001. Zhurn. 3: 179-180 (in Russian).
Nava, J.A., Jr. 1970. The reproductive biology of the364

References-,

Lawson, A.B., Henley, S.R. and G. Avery. 1986. Preyof leopards in four mountainous areas of the southwestern Cape Province. S. Afr. J. Wildl. Res. 16(2): 4752.
Noss, R.F. 1987. Corridors in real landscapes: a reply to
Simberloff and Cox. Conserv. Biol. l(2): 159- 164.
- . 199 1. Landscape connectivity: different functions at
different scales. Pp. 27-39 in W.E. Hudson, ed.
Landscape linkages and biodiversity. Island Press andDefenders of Wildlife, Washington, D.C.
Novkov, E. and Hanzl, R. 1968. [Contribution to knowledge of the role played by the lynx in forest communities.] Schweiz. 2. Forstwes.
1 19(2): 1 14- 126 (inFrench).
Novikov, G.A. 1962. Carnivorous mammals of the faunaof the U.S.S.R. Israel Program for Scientific
Translations, Jerusalem.
Nowak, R.M. 1976. The cougar in the United States andCanada. Unpubl. report, New York Zoological Society
and U.S. Fish and Wildlife Service, Washington, D.C.
- and Paradiso, J.L. 1983. Walker’s mammals of theworld, Vol. II. John Hopkins Univ. Press, Baltimore.
Nowell, K. 1990. Formosa and the clouded leopard. Cat
News 13: 15. Bougy-Villars, Switzerland.
- . 1991. Aboriginal hunting in the Tawu Mountain
Nature Reserve Area, Taiwan. Unpubl. report to theWildlife
Ecology Laboratory,
National Taiwan
University, Taipei.
- . 1993a. Tiger bone in Taipei. TRAFFIC Bulletin 13(3):112-l 14.
- . 1993b. Tiger bone medicines and trade. Cat News 18:
6-9. Bougy-Villars, Switzerland.
- 7Pei Chia-Jai and Chyi Wei-Lien. 1992. The horns of adilemma: Taiwan ‘s market for rhino horn. TRAFFIC
International, Cambridge, U.K.
- 9 Scheepers, L. and C. Brain. 1994. History of a maneater. Cat News 20: 9- 10.
Nuryatdinov, T. and Reimov, R. 1972. [Ecology of themost important mammals and birds of Karakalpakia.]
Tashkent(in Russian).
O’Brien, S.J. and Evermann, J.F. 1988. Interactive influence of infectious diseaseand genetic diversity in natural populations.TrendsEcol. Evol. 3( 10): 254-259.
- and Mayr, E. 1991. Bureaucratic mischief: recognizing endangeredspeciesand subspecies.Science 25 1:
1187-l 188.
-, Wildt, D.E., Goldman, D., Merril, C.R. and Bush, M.
1983. The cheetahis depauperatein genetic variation.
Science221: 459-462.
-, Roelke,M.E., Marker, L., Newman,A., Winkler, C.A.,
Meltzer, D., Colly, L., Evermann, J.F., Bush, M. and
Wildt, D.E. 1985.Genetic basisfor speciesvulnerability in the cheetah.Science227: 1428-1434.
-, Wildt, D.E. andBush, M. 1986.The cheetahin genetic

peril. Sci. Am. 254(5): 68-76.
-, Wildt, D.E., Bush, M., Caro, T.M., FitzGibbon, C.,
Aggundey, I. and Leakey, R.E. 1987a. East African
cheetahs: Evidence for two population bottlenecks?
Proc. Natl. Acad. Sci. USA 84: 508-511.
-, S.J., Martenson, J.S., Packer, C., Herbs&L., de Vos,
V., Joslin, P., Ott-Joslin, J., Wildt, D.E. and Bush, M.
1987b. Biochemical genetic variation in geographic
isolatesof African and Asiatic lions. Nat. Geog. Res.
3( 1): 114-124.
-, Joslin, P., Smith, G.L. III, Wolfe, R., Schaffer, N.,
Heath, E., Ott-Joslin, J., Rawal, P.P., Bhattacharjee,
K.K., and Martenson, J.S. 1987~.Evidence for African
origins of foundersof the Asiatic lion SpeciesSurvival
Plan. Zoo Biol. 6: 99-l 16.
-, Collier, G.E., Beneveniste, R.E., Nash, W.G.,
Newman, A.K., Simonson,J.M., Eichelberger, M.A.,
Seal, U.S., Janssen,D., Bush, M. and D.E. Wildt.
19874.Setting the molecularclock in Felidae: the greatcats,Panthera.Pp. lo-27 in R.L. Tilson and U.S. Seal,
eds. Tigers of the world. Noyes, Park Ridge, New
Jersey.
-, Roelke, M.E., Yuhki, N., Richards, K.W., Johnson,
W.E., Franklin, W.L., Anderson, A.E., Bass,O.L., Jr.,
Belden, R.C. and J.S. Martenson. 1990. Genetic introgression within the Florida panther Felis concoZor
coryi. Nat. Geog. Res.6(4): 485-494.
O’Connor, R.M. 1984. Population trends, age structure,and reproductive characteristics of female lynx in
Alaska, 1961 through 1973. M.S. thesis, Univ. of
Alaska, Fairbanks.
Ognev, S.I. 1926. A new genusand speciesof cat fromthe Transcaspian region. Ann. MUX 2001. Acad. Sci
U.S.S.R.27: 356-362.
- . 1935. Mammals of the U.S.S.R. and adjacent countries. Israel Program for Scientific Translations,
Jerusalem( 1962).
Olbricht, G. andSchiirer, U., eds.1993.International studbook for the black-footed cat, 1992. Zoologischer
Garten der Stadt Wuppertal, Wuppertal, Germany.
Oldeman, L.R., van Engelen, V.W.P. and J.H.M. Pulles.
1990. The extent of human-inducedsoil degradation.
Annex 5 of L.R. Oldeman,R.T.A. Hakkeling andW.G.
Sombroek, eds. World map of. the status ofI humaninducedsoil degradation, rev. 2d edn.International Soil
Reference and Information Center Wageningen,
Netherlands.
Oli, M.K. 1991.The ecology and conservationof the snow
leopard (Panthera uncia) in the Annapurna
Conservation Area, Nepal. M.Phil. thesis, Univ.
Edinburgh, Edinburgh.
- . 1994.Snow leopardsand a local humanpopulation ina protected area: a case study from the Nepalese
Himalaya. In J.L. Fox and Du Jizeng, eds.Proc. of* the365References

seventh intl. snow leopard symp. International Snow
Leopard Trust, Seattle.
- Taylor, I.R. and Rogers, M.E. 1994. Snow leopard‘(P an th era uncia) predation of livestock: an assessmentof local perceptions in the Annapurna Conservation
Area, Nepal. Biol. Conserv. 68: 63-68.
Oliveira, T.G. 1994. Neotropical Cats: Ecology andConservation. EDUFMA (Univ. Fed. do Maranhao),
San Luis, Brazil.
Olmsted, R.A., Langley, R., Roelke, M.E., Goeken, R.M.,
Adger-Johnson, D., Goff, J.P., Albert, J.P., Packer, C.,
Caro, T.M., Scheepers, L., Wildt, D.E., Bush, M.,
Martenson, J.S. and S.J. O’Brien. 1992. Worldwide
prevalence of lentivirus infection in wild Felidae
species: epidemiologic and phylogenetic aspects. J.
Viral. 66( 10): 6008-6018.
Olney, P.J.S. and Ellis, P., eds. 1991. Species of mammals bred in captivity during 1988/1989 and multiple
generation births. Int. Zoo Year-b. 30: 420-490.
Olrog, C.C. and Lucero, M.M. 1981. [Guide to the mammals of Argentina.] Fundacion Miguel Lillo, Tucuman(in Spanish).
Olson, J.S., Watts, J.A. and L.J. Allison. 1983. Carbon inlive vegetation ofmajor world ecosystems. U.S. Dept.
Energy publ. no. 1997, Envl. Sci, Div., Oak Ridge Natl.
Laboratory, Tennessee.
Osborn, D. and Helmy, I. 1980. The contemporary landmammals of Egypt (including Sinai). Fieldiana Zool.,
New Series 5: l-579.
Osborn, H,F. and Anthony, H.E. 1922. Can we save themammals? Nat. Hist. 22: 389-415.
Osgood, W.H. 1943. The mammals of Chile. Field. Mus.
Nat. Hist., 2001. Ser. 30: l-268.
Owens, M. and Owens, D. 1980. The fences of death. Afr.
Wildl. 4(6): 25-27.
- . 1984. Kalahari lions break the rules. Int. Wildl. 14: 413.
Oza, G.M. 1983. Deteriorating habitat and prospects of the
Asiatic lion. Environ. Cons. 6(3): 349-352.
Packer, C. 1986. The ecology of sociality in felids. Pp.
429-45 1 in D.I. Rubenstein and R.W. Wrangham, eds.
Ecological aspects of social evolution. Princeton Univ.
Press, Princeton.
- . 1990. Serengeti lion survey. In S.A. Huish and K.L.I.
Campbell, eds. Serengeti Wildlife Research Centre
biennial report of scientific results for 1988 and 1989.
Arusha, Tanzania.
- . 1992. Captives in the wild. Nat. Geog. April: 122-136.
- and Pusey, A.E. 1982. Cooperation and competition
within coalitions of male lions: kin selection or gametheory? Nature 296: 740-742.
- . 1983. Adaptations of female lions to infanticide byincoming males. Am. Nat. 121: 716-728.
-. 1987. Intrasexual cooperation and the sex ratio in

African lions. Am. Nat. 130: 636-642.
- . and Ruttan, L. 1988. The evolution of cooperative
hunting. Am. Nat. 132: 159-198.
-, Herbst, L., Pusey, A.E., Bygott, J.D., Hanby, J.P.,
Cairns, S.J. and M. Borgerhoff-Mulder.
1988.
Reproductive success of lions. Pp. 363-383 in T.H.
Clutton-Brock,ed. Reproductive
success. Univ.
Chicago Press, Chicago.
-, Scheel, D., and A.E. Pusey. 1990. Why lions form
groups: food is not enough. Am. Nat. 136: l-19.
-, Gilbert, D.A., Pusey, A.E. and S.J. O’Brien. 1991a. A
molecular genetic analysis of kinship and cooperationin African lions. Nature 35 1: 562-565.
-, Pusey, A.E., Rowley, H., Gilbert, D.A., Martenson, J.
and S.J. O’Brien. 1991b. Case study of a population
bottleneck: lions of the Ngorongoro Crater. Conserv.
Biol. 5(2): 219-230.
Paintiff, J.A. and Anderson, D.E. 1980. Breeding the margay at New Orleans Zoo. Int. Zoo. Yearb. 20: 223-224.
Pall, O., Jalkotzy, M. and I. Ross. 1988. The cougar inAlberta. Unpubl. report to Alberta Forestry, Lands andWildlife, Associated Resources Consultants, Calgary.
Palma, L.A. 1980. [On the distribution, ecology and conservation of the Iberian lynx in Portugal.] Actas I
Reunion Iberoamer. 2001. Vert. : 569-586. Sevilla,
Ministerio de Universidadese Investigation(inPortugese).
Palmer, R. and N. Fairall. 1988. Caracal and African wild
cat diet in the Karoo National Park and the implications
thereof for hyrax. S. Afr. J. WiZdZ.Res. 18: 30-34.
Paloheimo, J.E. and Fraser, D. 198 1. Estimation of harvest
rate and vulnerability from age and sex data. J. Wildl.
Manage. 45:948-958.
Palomares, F., Rodtiguez, A., Laffitte, R. and M. Delibes.
199 1. The status and distribution of the Iberian lynx,
Felis pardina (Temminck), in the Coto Dofiana area,
SW Spain. Biol. Conserv. 57: 159-169.
Palvaniyazov, M. 1990. [Mammals of the southern CisAral territory under conditions of anthropogenic
change as exemplified by carnivores.] Tashkent (inRussian).
Panwar, H.S. 1979. A note on tiger census technique basedon pugmark tracings. Indian Forester (Special Issue,Feb.): 70-77.
-. 1987. Project Tiger: the reserves, the tigers and their
future. Pp. 110-l 17 in R.L. Tilson and U.S. Seal, eds.
Tigers of the world: the biology, biopolitics, management, and conservation of an endangered species.
Noyes, Park Ridge, New Jersey.
- and Gopal, R. 1984. Conservation of wild cats in India.
Pp. 63-80 in The plight of the cats: proc. meeting andworkshop of the IUCN/SSC Cat Specialist Group at
Kanha National Park, Madhya Pradesh, India, 9-12
April 1984. Unpubl. report, IUCN/SSC Cat Specialist

366

References

Group, Bougy, Switzerland.
Parent, G.H. 1974. [Plea for the wildcat, or 10 excellent
reasons for protecting this misunderstood animal in
Belgium...and elsewhere] L’Homme et la Nature 10:
1- 15 (in French).
- . 1975. [Recent colonization of the wildcat Felis silvestris silvestris
Schreber in Belgian Lorraine.]
Mammalia 39: 251-288 (in French).
Parker, G.R. 198 1. Winter habitat use and hunting activities of lynx (Lynx canadensis) on Cape Breton Island,
Nova Scotia. Pp. 221-248 in J. Chapman and D.
Pursley, eds. Proc. worldwide
furbearer
conf
Frostburg, Maryland.
-, Maxwell, J.W., Morton, L.D., and Smith, G.E.J. 1983.
The ecology of the lynx (Lynx canadensis) on Cape
Breton Island. Can. J. 2001. 61: 770-786.
- and Smith, G.E.J. 1983. Sex- and age-specific reproductive and physical parameters of the bobcat (Lynxrufus) on Cape Breton Island, Nova Scotia. Can. J.
2001. 61: 1771-1782.
Paterson, H.E.H. 1985. The recognition concept of
species. Pp. 21-29 in E.S. Vrba, ed. Species andSpeciation. Transvaal Mus. Monograph no. 4, Pretoria.
Pathak, B.J. 1990. Rusty-spotted cat Felis rubiginosa
Geoffroy: a new record for Gir Wildlife Sanctuary andNational Park. J. Bombay Nat. Hist. Sot. 87: 445.
Patterson, J.H. 1907. The maneaters of Tsavo. Fontana,
London (1973).
Payne, J., Francis, C.M. and K. Phillips. 1985. A .field
guide to the mammals ofBorneo. The Sabah Society,
Malaysia.
Paz y Mifio, G. 1988. Notas sobre la caceria y la conservacion de 10s felidos en la Amazonia Ecuatoriana.
[Notes on the hunting and conservation of cat species in
Ecuador’s Amazonia.] Fundacidn Simon Bolivar
Boletin Cientzfico 2(3), Quito, Ecuador: 1-14.
Pearson, O.P. 1957. Additions to the mammalian fauna of
Peru and notes on some other Peruvian mammals.
Breviora 73: 1-7.
- . 1983. Characteristics
of a mammalian fauna fromforests in Patagonia, southern Argentina. J. Mamm.
64(3): 476-492.
Pennycuick, C.J. and Rudnai, J. 1970. A method of identifying individual lions Panthera leo with an analysis of
the reliability of identification. J. Zool., Lond. 160: 497508.
Perry, R. 1964. The world of the tiger. Cassell,London.
Peters,G. 1980.The vocal repertoire of the snow leopard(Uncia uncia, Schreber 1775).Intl. Ped. Book of Snow
Leopards2: 137-158.
Petersen,M.K. 1977a.Behaviour of the margay. Pp. 69-76
in R.L. Eaton, ed. The world’s cats 3(2). Carnivore Res.
Inst., Univ. Washington,Seattle.
- . 1977b.Courtship andmating patternsof margays.Pp.

22-35 in R.L. Eaton, ed. The world’s cats 3(3).
Carnivore Res. Inst., Univ. Washington, Seattle.
- and Petersen, M.K. 1978. Growth rates and other postnatal developmental changes in margays. Carnivore
l(1): 87-92.
Peterson, R.L. and Downing, S.C. 1952. Notes on the bobcat (Lynx rufus) of eastern North America with thedescription of a new race. Contr. Royal Ontario Mus.
33: l-23.
Peterson, R.O. and Page, R.E. 1988. The rise and fall ofIsle Royal wolves, 1975-1986. J. Mamm. 69: 89-99.
Pettifer, H. 198 1. The experimental release of captive-bredcheetah into the natural environment. Proc. Worldwide
Furbearer Conf 1: 1121-1142.
Petzsch, H. 1968. [The cats, 2d edn.] Urania, Leipzig (inGerman).
- . 1970. Kritisches Uber die neuentdeckte IriomoteWildkatze (Mayailurus iriomotensis). [Critical comments on the newly-discovered
Iriomote cat.] Das
Pelzgewerbe 20(5).
Pham Trong Anh. 1982. [Study of the Carnivora in thenorthern provinces of Vietnam.] Ph.D. thesis, Univ.
Hanoi, Hanoi (in Vietnamese).
Phillips, W.W.A. 1935. Manual of the mammalsof Sri
Lanka, 2d rev.edn: Part III. Wildlife and Nature
Protection Society of Sri Lanka, Columbo (1984).
Pickett, S.T.A., Parker, V.T. and P.L. Fiedler. 1992.The
new paradigmin ecology: implicationsfor conservation
biology above the specieslevel. Pp. 65-88 in P.L.
Fiedler and S.K. Jain, eds. Conservation biology: thetheory and practice of nature conservation,preservation and management.Chapmanand Hall, London.
Piechoki, R. 1986. [Distribution, losses,weights andmeasurementsof the wildcat (Felis silvestris Schreber,1777).] Hercynia 25: 235-258 (in German).
Pikunov, D.G. 1976. [Biology of the Amur leopard.] M.S.
thesis,Univ. Moscow, Moscow (in Russian).
- . 1988. Amur tiger (P.t. altaica): presentsituation andperspectives for preservation of its population in theSoviet Far East. Pp. 175-184 in B. Dresser,R. Reece
and E. Maruska, eds.Proc. 5th world con& on breeding endangeredspeciesin captivity. Cincinnati, Ohio.
- . 1994. Proposalsfor conserving the Amur tiger in the
RussianFar East. Cat News 20: 4-6. Bougy-Villars,
Switzerland.
- andKorkishko, V.G. 1989.The presentdistribution andnumbersof leopardsPanthera pardus in the Soviet Far
East. Transl. to Engl. reprinted in A. Shoemaker,ed.
1988 Internatl. leopard studbook. Riverbanks
Zoological Park, Columbia, South Carolina.
Pine, R.H., Bishop, I.R. and Jackson, R.J. 1970.
Preliminary list of mammals of the Xavantinal
Cachimboexpedition (Central Brazil). Transact.Royal
Sot. Tropical Med. and Hygiene 64: 668-670.

367

References

and felids. Proc. ann. meeting Sot. Theriogenology
1991: 334-337.
-, Gelwicks, E.J., Wachs, K.B., Keller, G.L., Maruska,
E.J. and B.L. Dresser. 1989. Successful interspecies
transferof embryosfrom the Indian desertcat (Felis silvestris ornata) to the domesticcat (FeZiscatus)following in vitro fertilization. Biol. Reprod. 4O(Suppl. 1):
61.
-, Keller, G.L. and B.L. Dresser.1993.In vitro fertilization in domestic and non-domestic cats, including
sequencesof early nuclear events in in vitro development, cryopreservation and sucessfulintra- and interspeciesembryo transfer. J. Reprod. Fert. Suppl. 47:
189-201.
Powell, J.R. 1974.Protein variation in natural populationsof animals.Evol. Biol. 8: 79-l 19.
Prater, S.H. 1971. The book of Indian mammals,3d edn.
Bombay Natural History Society, Bombay.
Prator, T., Thomas,W.D., Jones,M. and M. Dee. 1988.A
twenty-year overview of selected rare carnivores in
captivity. Pp. 191-229 in B. Dresser,R. Reeceand E.
Maruska, eds.Proc. 5th world con&on breedingendangered speciesin captivity. Cincinnati, Ohio.
Pringle, J.A. and V.L. Pringle. 1979. Observationson thelynx Felis caracal in the Bedford district. S. Afr. J.
Zool. 14: l-4.
Prins, H.H.T. and Iason, G.R. 1989. Dangerouslions andnonchalantbuffalo. Behaviour: 262-296.
Prynn, D. 1980. Tigers and leopardsin Russia’sFar East.
Oryx 15: 496-503.
Pulliainen,E. 1981. Winter diet of Felis Lynxin SEFinland
ascomparedwith the nutrition of other northern lynxes.
2. Saugetierk.46(4): 249-259.
- . 1992. From extinction to real lynx life: Finnish experiences.Pp. 17-18 in The situation, conservationneeds
and reintroduction of lynx in Europe. Proc. symp. 1719 October, Neuchatel.Council of Europe, Strasbourg.
-, Lindgren, E. and P.S. Tunkkari. 1988. [Diet of lynx inwinter: hare or white-tailed deer?] Jagaren 3: 42-44(in Swedish).
Pusey, A.E. and Packer, C. 1987. The evolution of sexbiaseddispersalin lions. Behaviour 101: 275-310.
Puzachenko, A.Yu. 1993a.The European wildcat (Felis
silvestris) in Armenia, Azerbaijan, Belarus, Georgia,
Moldova, RussiaandUkraine. Pp. 64-71 in Proc. seminar on the biology and conservation of the wildcat(Felis silvestris), Nancy, France, 23-25 September
1992. Council of Europe, Strasbourg.
- . 1993b. On the taxonomic status of Felis silvestris
Schreber, 1777, in the Caucasuswith somecommentson the variation between the European and African
wildcat. Pp. 75-81 in Proc. seminaron the biology andconservation of the wildcat (Felis silvestris), Nancy,
France, 23-25 September 1992. Council of Europe,-,

Miller, S.D. and M.L. Schamberger.
1979.
Contributions to the mammalogy of Chile. Mammalia
43(3): 339-376.
Pocock, R.I. 1907a. Notes upon some African species of
the genus Fe&s, based upon specimens recently exhibited in the Society’s Gardens. Proc. 2001. Sot. Lond.
1907: 656-677.
- . 1907b. On Pallas’s cat. Proc. 2001. Sot. Lond. 1907:
299-300.
-. 1907~. On English domestic cats. Proc. Zool. Sot.
Lond. 1907: 143-168.
- . 19 16. On some of the cranial and external characters of
the hunting leopard or cheetah. Ann. Mag. Nat. Hist.
8(18): 419-429.
-. 19 17. The classification of existing Felidae. Ann. Mag.
Nat. Hist. 8(20).
- . 1927. Description of a new species of cheetah(Acinonyx). Proc. 2001. Sot. London 1927: 245-25 1.
- . 1930. The panthers and ounces of Asia. J. Bombay Nat.
Hist. Sot. 1: 63-82; 307-336.
- . 1932a. The leopards of Africa. Proc. 2001. Sot., Lond.:543-59 1.
- . 1932b. The marbled cat (Pardofelis marmorata) andsome other Oriental species, with the definition of a
new Genus of the Felidae. Proc. 2001. Sot. Lond. 1932:
741-766.
-. 1938. A new race of the sand cat (Felis margarita).
Ann. Mag. Nat. Hist. 11: 472-476.
- . 1939a. The fauna of British India, Mammalin, I.
Primates and Carnivora, 2d edn. Taylor and Francis,
London.
- . 1939b. The races of the jaguar, Panthera onca. Novit.
Zool., Tring. 4 1: 406-422.
-. I94 1. Thefauna of British India, Mammalia, II Taylor
and Francis, London.
-. 1951. Catalogue of the Genus Felis. British Museumof Natural History, London.
Poland, H. 1892. Fur-bearing animals in nature and incommerce. Gurney and Jackson, London.
Poland, J.D., Barnes, A.M. and J.J. Herman. 1973. Humanbubonic plague from exposure to naturally infectedwild carnivore. Am. J. Epidemiol. 97: 332-337.
Pollack, E.M. 1950. Breeding habits of the bobcat in northeastern United States. J. Mamm. 31: 327-330.
Polo, Marco. 1299. Travels of Marco Polo. Dent, London(1954).
Poole, K.G. 1992. Lynx research in the Northwest
Territories, 1991-92. Unpubl. report, Government of
the Northwest Territories, Yellowknife.
- . 1994.Characteristicsof an unharvestedlynx population during a snowshoehare decline.J. Wildl. Manage.
58 (4): 608-618.
Pope, C.E. and Dresser, B.L. 1991. Development ofassistedreproductiontechniquesin nondomesticbovids

368References

Strasbourg.
Quigley, H.B. and Crawshaw, P.G. 1992. A conservation
plan for the jaguar Panthera onca in the Pantanalregion of Brazil. Biol. Conserv. 6 1: 149- 157.
- and Homocker, M.G. 1992. Large carnivore ecology:
from where do we come and to where shall we go? Pp.
1089- 1097 in D.R. McCullough and R.H. Barrett, eds.
Wildlife 2001: populations. Elsevier, New York andLondon.
Quillen, P. 1992. Cooperative captive breeding initiatives
for small cat species. Pp. 33 in D.E. Wildt, J.D. Mellenand U.S. Seal, eds. 1992. Felid action plan, 1991 and1992: AAZPA Felid Taxon Advisory Group regional
collection plan and IUCN Captive Breeding Specialist
Group global felid captive action plan. National
Zoological Park, Front Royal, Virginia.
Quinn, N.W.S. and Gardner, J.F. 1984. Relationships ofage and sex to lynx pelt characteristics.
J. WiZdZ.
Manage. 48: 953-956.
- and Parker, G. 1987. Lynx. Pp. 683-694 in M. Novak,
J. Baker, M. Obbard, and B. Malloch, eds. Wild
furbearer management and conservation in North
America. Ontario Ministry of Natural Resources,
Toronto.
- and Thompson, J.E. 1985. Age and sex of trapped lynx,
Felis canadensis, related to period of capture and trapping technique. Can. Field-Nat. 99: 267-269.
Rabinovich, J., Capurro, A., Folgarait, P., Kitzberger, T.,
Kramer, G., Novaro, A., Puppo, M. and A. Travaini.
1987. [State of knowledgeof12 speciesofforestfaunaof commercial value in Argentina.] Unpubl. report,
IUCN, BuenosAires (in Spanish).
Rabinowitz, A.R. 1986.Jaguarpredationon domesticlivestock in Belize. Wildl. Sot. Bull. 14: 170-174.
- . 1988.The cloudedleopardin Taiwan. O~X 22( 1): 4647.
- . 1989. The density and behavior of large cats in a drytropical forest mosaic in Huai Kha Khaeng Wildlife
Sanctuary, Thailand. Nat. Hist. Bull. Siam. Sot. 37(2):235-251.
- . 1990.Noteson the behavior andmovementsof leopard
cats,Felis bengalensis,in a dry tropical forest mosaicin
Thailand. Biotropica 22(4): 397-403.
- . 199I. Belize trip report, June 1%July 30 1991.
Unpubl. report, Wildlife Conservation Society, Bronx,
New York. Summarized in Cat News 16: 9. BougyVillars, Switzerland.
- . 1992. Chasingthe dragon’s tail: the struggle to saveThailand’swild cats. Doubleday, New York.
- . 1993.Estimating the Indochinesetiger Panthera tigris
corbetti populationin Thailand. Biol. Conserv.65: 213217.
- andNottingham, B.G. 1986.Ecology andbehaviour of
the jaguar (Panthera onca) in Belize, Central America.

J. Zool., Lund. 210: 149-159.
- andWalker, S.R. 1991.The carnivore community in a
dry tropical forest mosaicin Huai Kha KhaengWildlife
Sanctuary, Thailand. J. Troop.Ecol. 7: 37-47.
-, Andau, P. and P.P.K. Chai. 1987. The clouded leopard in Malaysian Borneo. Ory.~22(2): 107-111.
Rabor, S. 1986. Guide to Philippineflora andfauna II:
birds and mammaIs.Natural ResourcesManagement
Center, Ministry of Natural Resources and Univ.
Philippines.Manila.
Ragni, B. 1978. Observationson the ecology and behaviour of the wild cat (Felis silvestris Schreber, 1777) in
Italy. Carniv. Genet. Newsl. 3: 270-274.
- . 1981. [Forest wildcat, FeZissilvestrisSchreber, 1777.1
Pp. 105-l 13 in [Distribution and biology of22 mammalian speciesin Italy.] C.N.R., Rome (in Italian).
- 1988. [Status and conservation problemsof the felids‘(Felidae) in Italy.] Supl. Ric. Biol. Selv. 14: 455-477(in Italian).
- . 1993a.Statusand conservationof the wildcat in Italy.
Pp. 40-41 in Proc. seminaron the biology and conservation of the wildcat (Felis silvestris), Nancy, France,
23-25 September1992. Council of Europe, Strasbourg.
- . 1993b. The crucial problem of in vivo identificationof wildcat and recognition of hybrids with domestic
cats. Pp. 86-88 in Proc. seminar on the biology andconservation of the wildcat (Felis silvestris), Nancy,
France, 23-25 September 1992. Council of Europe,
Strasbourg.
- 9Possenti,M., Guidali, F., Mingozzi, T. and G. Tosi.
1992. Statusof the lynx in Italy. Pp. 74-76 in The situation, conservationneedsand reintroduction of lynx inEurope. Proc. symp. 17-l 9 October, Neuchatel.
Council of Europe; Strasbourg.
Rahm,U. 1966.[Mammals of the equatorialforest of eastern Congo.] Ann. Mus. afr. centr., Tervuren, SC.2001.
(in French).
Rahm,U. andChristiaensen,A. 1963.Les Mammiferesdela region occidentale du lac Kivu. [Mammals of thewestern region of Lake Kivu.] Ann. Mus. afr. centr.,
Tervuren, SC.Zoo1149: 39-121.
Rails, K. and Ballou, J.D. 1983. Extinction lessonsfrom
zoos. Pp. 164-l 84 in C.M. Schonewald-Cox, S.M.
Chambers, B. MacBryde and W.L. Thomas, eds.
Geneticsand conservation: a referencefor managing
wild animal and plant populations. BenjaminCummings,Menlo Park, California.
- 9and A. Templeton. 1988. Estimatesof lethal equivalentsand the costsof inbreedingin mammals.Conserv.
Biol. 2: 185-193.
Randi, E. and Ragni, B. 1986. Primi resultati dell’analisi
elettroforetica di un campionedi populazioni italiane digatto selvatico e di gatto domestico. [First results ofthe electrophoreticanalysisof a sampleof Italian popu-369

References

mammals.WiZdZ.Sot. Bull. 14: 255261.
Regan, T. and Maehr, D. 1990. Melanistic bobcats in
Florida. Florida Field Nat. 18(4): 84-87.
Rengger,J. R. 1830. [Natural history of the mammalsof
Paraguay.] Base1(in German).
Reza Khan, Mohd.A. 1986.The statusand distribution of
the cats in Bangladesh. Pp. 43-49 in S.D. Miller and
D.D. Everett, eds. Cats ofthe world: biology, conservation, and management.National Wildlife Federation,
Washington,D.C.
Rice, C. 1986. Observations of predators and prey in
Eravikulam National Park, Kerala. J. Bombay Nat.
Hist. Sot. 83(2): 283-305.
Rieger, I. 1984. Oestroustiming in ounces,Uncia uncia,
Schreber(1775). Intl. Ped. Book ofSnow Leopards4:
99-103.
Riols, C. 1988. [Study of the diet of the forest wildcat(Felis silvestris)in easternFrance.] Bull. Mens. 0. N. C.
121: 22-27 (in French).
Rishi, V. 1988. Man, mask and man-eater. Tigerpaper
15(3): 9- 14.
Roberts, A. 1951. The mammals of South Africa.
Mammals of South Africa Book Fund, Johannesburg.
Roberts, T.J. 1977. The mammalsof Pakistan. Ernest
Benn, London.
Robinette, W.I., Gashwiler, J.S. and O.W. Morris. 1961.
Noteson cougarproductivity andlife history. J. Mamm.
42: 204-217.
Robinson,I.H. and Delibes,M. 1988. The distribution of
faeces by the Spanish lynx (Felis pardina). J. Zool.,
Lond. 2 16: 577-582.
Robinson, R. 1976. Homologousgenetic variation in the
Felidae. Genetica 46: l-3 1.
- . 1991.Geneticsfor cat breeders.Pergamon,Oxford.
Rodenberg, W.F. 1977. The trade in wild animal furs in
Afghanistan. Unpubl. report to UNDP/FAO, Kabul.
Rodgers,W.A. 1974.The lion (Panthera leo, Linn.) population of the easternSelous Game Reserve. E. Afr.
Wild/. J. 12: 313-317.
Rodriguez, A. and Delibes, M. 1990. [The Iberian lynx+
Lynx pardina in Spain: distribution and conservation
problems.] ICONA, Madrid (in Spanish).
- . 1992.Current rangeandstatusof the Iberian lynx Felis
pardina Temminck, 1824in Spain. Biol. Conserv. 61:
189-196.
Rodriguez, F. and Paz y Mifio, C.G. 1989. [Fe/id collections represented in the principal Ecuador-inn museums.] Unpubl. report, IUCN, Gland (in Spanish).
Roelke,M.E., Martenson, J.S. and S.J. O’Brien. 1993.The
consequencesof demographic reduction and genetic
depletionin the endangeredFlorida panther.Curr. Biol.
3: 340-350.
Rolley, R.E. 1983.Behavior and population dynamicsof
bobcats in Oklahoma. Ph.D. thesis, Oklahoma State

lations of wildcat and domestic cat.] I1 Convegnodell ‘Unione 2001. Italiana. Rome, Italy (in Italian).
- . 199 1. Genetic variability and biochemical systematics
of domestic and wild cat populations (Felis silvestris:
Felidae). J. Mamm. 72: 79-88.
Rasheed, S. and Gardner, M.B. 198 1. Isolation of feline
leukemia virus from a leopard cat cell line and search
for retrovirus in wild Felidae. J. Nat. Cancer Inst. 67:
929-933.
Rashid, M.A. 1984. Notes on conservation of the Asiatic
lion. Pp. 11 l- 114 in P. Jackson ed. The plight of thecats: proc. meeting and workshop of the IUCN/SSC Cat
Specialist Group at Kanha National Park, Madhya
Pradesh, India, 9-12 April 1984. Unpubl. report,
IUCN/SSC Cat Specialist Group, Bougy-Villars,
Switzerland.
- . 1991. Asiatic lion population up. Cat News 13: 12.
Bougy-Villars, Switzerland.
- and David, R.B. 1992. The Asiatic lion. Dept. of theEnvironment, Government of India, Baroda.
Rathore, F.S. and Thapar, V. 1984. Behavioral observations of leopard and jungle cat in Ranthambhor
National Park and Tiger Reserve,Rajasthan.Pp. 136139 in The plight of the cats: proc. meetingand workshopof the IUCN/SSC Cat SpecialistGroup at Kanha
National Park, Madhya Pradesh, India, 9-12 April
1984.Unpubl. report, IUCN/SSC Cat SpecialistGroup,
Bougy-Villars, Switzerland.
Rautenbach, I.L. 1978. The mammalsof the Transvaal.
Ph.D. thesis,Univ. of Natal, Pietermaritzburg.
Ravi Chellam. 1987. Asiatic lion study. Cat News6: 3 1.
Bougy-Villars, Switzerland.
- . 1993. Ecology of the Asiatic lion (Panthera leo persica).Ph.D. thesis,SaurashtraUniversity, Rajkot.
- and Johnsingh,A.J.T. 1993a.Management of Asiatic
lions in the Gir Forest, India. In N. DunstoneandM.L.
Gorman, eds. Mammals as predators. Proc. Symp.
Zool. Sot. Lond. 65. Clarendon,Oxford.
- . 1993b.Asiatic Zion(Pantheraleo persica).Unpubl. data
sheet,IUCN/SSC Cat SpecialistGroup, Bougy-Villars,
Switzerland.
Read, B. 1982. ChineseMateria Medica Vol. 4: animaldrugs. SouthernMaterials Center, Taipei.
Read,J.A. 1981. Geographicvariation in the bobcat (Felis
rufus) in the southcentral United States. M.S. thesis,
Texas A & M Univ., CollegeStation.
Redford, K.H. and Eisenberg,J.F. 1992.Mammals of the
Neotropics, Vol. 2: the southern cone. Univ. Chicago
Press,Chicago.
-, Taber, A. andJ. Simonetti. 1990.There is moreto biodiversity than tropical rain forests. Conserv.Biol. 4(3):328-330.
Reed, J.M., Doerr, P.D. and J.R. Walters. 1986.
Determining minimum population sizesfor birds and370

References

the number of subspecies. Bobcat Res. Conf Natl.
Wildl. Fed. Sci. Tech. Ser. 6: 80-86.
Sanderson, I.T. 1940. The mammals of the north
Cameroon forest area. Transactions 2002. Sot. London
24, pt. 7.
Sankhala, KS. 1967. Breeding behaviour of the tiger inRajasthan. Int. Zoo Yearb. 7: 133-147.
- . 1978. Tiger? The story of the Indian tiger. Collins,
London.
- . 1993. Return of the tiger. Lustre Press, New Delhi.
Santiapillai, C. and Ashby, K.R. 1988. The clouded leopard in Sumatra. &yx 22( 1): 44-45.
- 7Chambers, M.R. and N. Ishwaran. 1982. The leopard
Panthera pardus fusca (Meyer 1794) in the Ruhuna
National Park, Sri Lanka, and observations relevant to
its conservation. Biol. Conserv. 24: 5-14.
- and Ramono, W.S. 1992. Status of the leopard(Panthera pardus) in Java, Indonesia. Tigerpaper 19:
l-5.
- and Suprahman, H. 1985. On the status of the leopard
cat in Sumatra. Tigerpaper 12(3): 8- 13.
Sanyal, P. 1983. Mangrove tiger land: the Sundarbans ofIndia. Tigerpaper lO(3): l-4.
- . 1992. Catland around Calcutta. Tigerpaper 19: 18-20.
Sapozhenkov, Yu. F. 1960. [Caracal around Repetek.]
Priroda 2: 107-108 (in Russian).
- . 196 la. [Distribution and ecology of the sand cat Fehs
margarita in eastern Turkmenia.] 2002. Zh. 40: 10861089 (in Russian).
- . 1961b. [Ecology of the steppe cat Fehs Zybica Forst., in
the eastern Karakum.] ZooZZh. 40: 1585-1586 (inRussian).
- . 1962. [Ecology of the caracal (Felis caracal Mull.) in
the Karakum.] ZooZZh. 41: 11 lo- 1112 (in Russian).
Satunin, C. 1904. The black wild cat of Transcaucasia.
Proc. 2001. Sot. Lond. 1904: 163- 164.
Saunders, J.K. 1961. The biology of the Newfoundlandlynx. Ph.D. thesis, Cornell Univ., Ithaca.
- . 1963. Movements and activities of the lynx in
Newfoundland. J. Wildl. Manage. 27: 390-400.
Sausman, K. 1989. Sand cat, Felis margarita, international studbook, 31 December 1989. The Living Desert,
Palm Desert, California.
- 199 1. Sand cat, Felis margarita international studbook,
31 December 1991. The Living Desert, Palm Desert,
California.
Savage, R.J.G. 1978. Carnivores. Pp. 249-265 in V.J.
Maglio and H.B.S. Cooke, eds. Evolution of African
mammals. Harvard Univ. Press, London.
Sawarkar, V.B. 1986. Animal damage: predation on
domestic livestockby large carnivores.
IndianForester, October.
Sawyer, J. 1993. Plantations in the tropics: environmental concerns.
IUCN,
Gland, Switzerland
andUniv., Stillwater.
- . 1985. Dynamics of a harvested bobcat population in
Oklahoma. J. WildZ. Manage. 49: 283-292.
- . 1987. Bobcat. Pp. 670-681 in M. Novak, J. Baker, M.
Obbard, and B. Malloch eds. Wildfurbearer management and conservation in North America. Ontario
Ministry of Natural Resources, Toronto.
Rollings, C.T. 1945. Habits, food and parasites of the bobcat in Minnesota. J. Wildl. Manage. 9: 13 1- 145.
Rosevear, D.R. 1974. The carnivores of West Africa.
British Museum (Natural History), London.
Ross, P.I. and Jalkotzy, M.G. 1992. Characteristics of ahunted population of cougars in southwestern Alberta.
J. Wildl. Manage. 56(3): 417-426.
Rowe-Rowe, D.T. 1978. The small carnivores of Natal.
Lammergeyer 23: 22-27.
- . 1992. The carnivores of Natal. Natal Parks Board,
Pietermaritzburg, South Africa.
Rudnai, J. 1973. The social life of the lion. Medical andTechnical Publishing, Lancaster, U.K.
- . 1974. The pattern of lion predation in Nairobi Park. E.
Afr. Wildl. J. 12: 213-225.
Ruggiero, R.G. 1991. Prey selection of the lion (Panthera
Zeo L.) in the Manovo-Gounda-St. Floris National Park,
Central African Republic. Mammalia 55: 23-33.
Rust, W.D. 1980. Scent station transects as a means ofindexing bobcat population fluctuations. M.S. thesis,
Tennessee Tech. Univ., Cookeville.
Ryder, O.A., Shaw, J.H. and C.M. Wemmer. 1988.
Species, subspecies and ex situ conservation. Int. Zoo.
Yearb. 27: 134- 140.
Saberwal, V. 1990. Lion-human conflict around Gir. Bull.
Wildlife Institute of India 5(3): 28-30.
- 7Gibbs, J.P., Chellam, R. and A.J.T. Johnsingh. 1994.
Lion-human conflict in the Gir Forest, India. Conserv.
Biol. 8(2): 501-507.
Sadleir, R.M.F.S. 1966. Notes on reproduction in thelarger Felidae. Int. Zoo Yearb. 6: 184- 187.
Sale, J.B. and Berkmiiller, K. 1988. Manual of wildlife
techniques for India. United Nations Food andAgriculture Organization, Dehra Dun, India.
Salikhbaev, Kh.S. 1950. [Fur-bearing animals of the AmuDar’ya Delta (Karakalpakia) and measures toward theirrational use.] In [Information on Productivity Strengths
in Uzbekistan I], Uzbek Acad. Sci. (in Russian).
Salles, L. 1992. Felid phylogenetics: extant taxa and skull
morphology. Amer. Mus. Novitates 3047: l-67.
Salter, R. 1983. Summary of currently available information on internationally threatened wildlife species in
Burma. Unpubl. report to the FAO, Rangoon.
- . 1993. Wildlife in Lao PDR: a status report. IUCN,
Vientiane.
Samson, F.B. 1979. Multivariate analysis of cranial characters among bobcats with a preliminary discussion ofl371

References

vestris Schreber 1777en Suisse.[The Europeanforest
wildcat in Switzerland.] Rev. Suisse2001.77: 127-160.
-. 1974.Donneesnouvelles sur le Chat dessablesFelis
margarita Loche, 1858. [New information on the sandcat.] Rev. Suisse2001.81: 949-969.
-. 1978. Note sur la reproduction du manul Otocolobus
manul (Pallas, 1776). [Note on the reproduction of themanul.] Mammalia 42(3): 355-358.
-. 1979.La reproductiondu chat desmarais,Felis chaus(Guldenstadt,1776). [Reproductionof the swampcat.]
Mammalia 43(2): 215-223.
Scheel, D. 1993. Profitability, encounter rates, and preychoice of African lions. Behav. Ecol. 4: 90-97.
Scheepers,J.L. and Gilchrist, D. 1991. Leopardpredation
on giraffe calvesin the EtoshaNational Park. Madoqua
18(l): 49.
Scheffel, W. and Hemmer, H. 1974.Notizen zur Haltung
und Zucht der Sandkatze (Felis margarita Loche1858).[Notes on the careand breedingof the sandcat.]
2001.Gart. N.F. 44: 338-348.
Schonewald-Cox, C., Azari, R., and S. Blume. 1991.
Scale, variable density, and conservationplanning for
mammaliancarnivores. Conserv.Biol. 5(4): 49 l-495.
Schotts, E.B. Jr., Andrews, C.L. and T.W. Harvey. 1975.
Leptospirosisin selectedwild mammalsof the Florida
panhandleand southwesternGeorgia.J. Am. Vet. Med.
Assoc. 167: 587-589.
Scott, D.M. and Poole, C.M. 1989. A status overview of
Asian wetlands.Publ. no. 53, Asian WetlandsBureau,
Kuala Lumpur.
Scott, M.E. 1988.The impact of infection and diseaseon
animalpopulations:implications for conservationbiology. Conserv.Biol. 2(l): 40-56.
Scott, R., Easterbee,N. and D. Jefferies. 1993. A radiotracking study of wildcats in westernScotland. Pp. 9497 in Proc. seminar on the biology and conservationof. the wildcat (Felis silvestris), Nancy, France, 23-25
September1992. Council of Europe, Strasbourg.
Scrocchi, G.J. and Halloy, S.P. 1986. [Systematic, ecological, ethological and biogeograhpical notes on the
Andean mountain cat Felisjacobita Cornalia (Felidae,
Carnivora).] Acta. 2001. Lilloana 38(2): 157-170 (inSpanish).
Seager,S. and Desmorest,C. 1978. Reproductionof captive wild carnivores. In M. Fowler, ed. Zoo and wild
animal medicine.Saunders,Philadelphia.
Seal,U.S. andFoose,T.J. 1992. 1992executive summary:conservationasessmentand managementplan. P 13 in
D.E. Wildt, J.D. Mellen and U.S. Seal,eds. 1992.Fehd
action plan, 1991 and 1992: AAZPA Felid Taxon
Advisory Group regional collection plan and IUCN
Captive Breeding SpecialistGroup globalfelid captive
action plan. National Zoological Park, Front Royal,
Virginia.

Cambridge, U.K.
Sayer, J. 1983. Nature conservation and national parks.
Final report, Forest Management Project, Lao People’s
Democratic Republic and UN FAO, Rome.
- and A.A. Green. 1984. The distribution and status oflarge mammals in Benin. Mammal Rev. 14(l): 37-50.
- and van der Zon, A.P.M. 1981. National parks andwildlife management-Afghanistan.
A contribution to aconservation
strategy. Unpubl. report, Govt. ofAfghanistan and UNDP/FAO, Rome.
-, Harcourt, C.S. and N.M. Collins. 1992. The conservation atlas of tropical forests: Africa. IUCN and
Macmillan, London.
Schaffer, N. and Rosenthal, M. 1984. Report on theflatheaded cat reproductive projects. Unpubl. report,
Lincoln Park Zoo, Chicago.
Schaffer, W.M. 1984. Stretching and folding in lynx fur
returns: evidence for a strange attractor in nature? Am.
Nat. 124: 798-820.
Schaller, G.B. 1967. The deer and the tiger. Univ. of
Chicago Press, Chicago.
-. 1968. Hunting behaviour of the cheetah in the
Serengeti National Park, Tanzania. E. Afr. Wildl. J. 6:
95-100.
- . 1972. The Serengeti lion. Univ. of Chicago Press,
Chicago.
-. 1976. Mountain mammals in Pakistan. Oryx 13: 35 l356.
- . 1977. Mountain monarchs: wild sheep and goats of theHimalaya. Univ. of ChicagoPress,Chicago.
-. 1983. Mammals and their biomass on a Brazilian
ranch. Arq. 2001.Sao Paulo 3 1: l-36.
-. 1990.Saving China’swildlife. Int. Wildl. 20(1): 30-40.
- and Crawshaw, P.G. 1980. Movement patterns ofjaguar. Biotropica 12: 161-168.
- and Vasconcelos, J.M.C. 1978. Jaguar predation on
capybara.2. Saugetierk.43: 296-301.
-, Hu Jinchu, Pan Wenshi and Zhu Jing. 1985.The giant
pandasof Wolong. Univ. of Chicago Press,Chicago.
- 7Li, H., Lu, H., Ren, J.R., Qiu, M.J. and H.B. Wang.
1987. Status of large mammals in the Taxkorgan
Reserve,Xinjiang, China. Biol. Conserv.42: 53-71.
-, Ren, J.R. and M.J. Qiu. 1988a.Statusof snowleopard(Panthera uncia) in Qinghai and Gansu provinces,
China. Biol. Conserv.45: 179-194.
-, Li, H.T., Ren, J.R. and M.J. Qiu. 1988b. The snow
leopardin Xinjiang, China. Oryx 22: 197-204.
-, Tserendeleg,J. and G. Amarsanaa.1994.Observations
on snow leopards in Mongolia. In J.L. Fox and Du
Jizeng, eds. Proc. of the seventh intl. snow leopard
symp.International Snow Leopard Trust, Seattle.
Schantz, V.S. 1939.A white-footed bobcat. J. Mamm. 20:
106.
Schauenberg,P. 1970.Le chat forestierd’Europe Felis sil-372

References

Sarawak. Sarawak Mus. J. 4(3): 263-266.
Senner, J.W. 1980. Inbreeding depression and the survivalof zoo populations. Pp. 209-224 in M.E. Soule and B.A.
Wilcox, eds. Conservation biology: an evolutionarySunderland,
ecologicalperspective.
Sinauer,
Massachusetts.
Serhal, A. 1985. [Mammals of Lebanon.] Society forProtection of Nature and Natural Resources in
Lebanon, Beirut (in Arabic).
Serpell, J.A. 1988. The domestication and history of thecat. Pp. 151-158 in D.C. Turner and P. Bateson, eds.
The domestic cat: the biology of its behaviour.
Cambridge Univ. Press, Cambridge.
- . 1992. Domestic cats. Pp. 184- 189 in J. Seidensticker
and S. Lumpkin, eds. Great cats. Merehurst, London.
Severtzov, M.N. 1857-58. Notice sur la classification multieriale des carnivores, specialement des felides, et lesetudes de zoologie g&&ale qui s’y rattachent. [Notes
on the classification of carnivores, especially the felids,
and associated general zoological studies.] Revue etMagazine de Zoologie 9: 389-391,433-439;
10: 3-8,
145-150, 192-199,241-246, 385-393.
Seymour, K.L. 1989. Panthera onca. Mammalian Species
340: l-9.
Shafer, C.L. 1990. Nature reserves: island theory and conservation practice. Smithsonian Institution Press,
Washington, D.C.
Shaffer, M.L. 1987. Minimum viable populations: copingwith uncertainty. Pp. 69-86 in M.E. Soul& ed. Viable
populations for conservation. Cambridge Univ. Press,
Cambridge.
Sharma, I.K. 1979. Habits, feeding, breeding and reaction
to man of the desert cat Felis Zibyca (Gray) in the IndianDesert. J. Bombay Nat. Hist. Sot. 76(3): 498-499.
Sharma, V. and Sankhala, K. 1984. Vanishing cats ofRajasthan. Pages 117-135 in The plight of the cats:proc. meeting and workshop of the IUCN/SSC Cat
Specialist Group at Kanha National Park, Madhya
Pradesh, India, 9-12 April 1984. Unpubl. report,
IUCN/SSC
Cat Specialist Group, Bougy-Villars,
Switzerland.
Shaw, H.G. 1977. Impact of mountain lion on mule deerand cattle in northwestern Arizona. Pp. 17-32 in R.I.
Phillips and C. Jonkel, eds. Proc. 1975predator symp.,_
Montana For. and Conserv. Exp. Stn., Univ. Montana,
Missoula.
- . 1989. Soul among lions. Johnson Books, Boulder,
Colorado.
Shchadinov, Y. 1989. Cruel to be kind: reducing the Amurtiger population in the Soviet Far East is the only way
its survival in the natural world can be guaranteed.
Soviet Weeklv 16 Nov: 7.
Shi, Y. 1983. [On the influence of rangeland vegetation to
the density of plateau pikas (Ochotona curzoniae).]-, Lacy, R.C., et al. 1989. Florida panther Felis concolor
coryi viability analysis and species survival plan.
IUCN/SSC Captive Breeding Specialist Group, Apple
Valley, Minnesota.
- 7 Ellis, S.A., Foose, T.J. and A.P. Byers. 1993.
Conservationassessment and management plans(CAMPS) and global captive action plans (GCAPs).
CBSG News 4(2): 5-10.
Seidel, B. and Wisser, J. 1987. Clinical diseases of captive tigers-European
literature. Pp. 205230 in R.L.
Tilson and U.S. Seal, eds. Tigers ofthe world: the biology, biopolitics, management, and conservation of anendangered species. Noyes, Park Ridge, New Jersey.
Seidensticker, J. 1976. On the ecological separation
between tigers and leopards. Biotropica 8: 225234.
- . 1983. Predation by Panthera cats and measures ofhuman influence in habitats of South Asian monkeys.
Inter. J. Primatol. 4(3): 323-326.
- . 1986. Large carnivores and the consequences of habitat insularization: ecology and conservation of tigers inIndonesia and Bangladesh. Pp. l-42 in S.D. Miller and
D.D. Everett, eds. Cats ofthe world: biology, conservation and management. National Wildlife Federation,
Washington, D.C.
- . 199 la. Leopards. Pp. 106- 115 in J. Seidensticker and
S. Lumpkin, eds. Great cats. Merehurst, London.
- . 1991b. Pumas. Pp. 130-137 in J. Seidensticker and S.
Lumpkin, eds. Great cats. Merehurst, London.
- and Lumpkin, S., eds. 1991. Great cats. Merehurst,
London.
- . 1992. Mountain lions don’t stalk people. True or false?
Smithsonian 22( 11): 113- 122.
Seidensticker, J. and McDougal, C. 1993. Tiger predatory
behaviour, ecology and conservation. In N. Dunstone
and M.L. Gorman, eds. Mammals as predators. Proc.
Symp. 2001. Sot. Lond. 65. Clarendon, Oxford.
-, Hornocker, M., Wiles, W. and Messick, J. 1973.
Mountain lion social organization in the Idaho
Primitive Area. WiZdZ.Monogr. 35: I-61.
- 7 Lahiri, J.E., Das, KC. and A. Wright. 1976. Problem
tiger in the Sunderbans. Oryx 11: 267-273.
-, Sunquist, M.E. and C. McDougal. 1990. Leopards living at the edge of the Royal Chitwan National Park,
Nepal. Pp. 415-423 in J.C. Daniel and J.S. Serrao, eds.
Conservation in developing countries: problems andprospects. Proc. Centenary Seminar of the Bombay
Nat. Hist. Sot. Bombay Nat. Hist. Sot. and Oxford
Univ. Press, Bombay.
Seifert, S. and Miiller, P. 1976. International tiger studbook. Zoologischer Garten Leipzig, Leipzig.
Selander, R.K. and Johnson, W.E. 1973. Genetic variation among vertebrate species. Annual Review of
Ecology and Systematics 4: 75-9 1.
Selous, E.M. and Banks, E. 1935. The clouded leopard in373

References

Carpathians. Biologia Bratisl. 28: 127- 137.
Sliwa, A. 1993. A habitat description and first data on
ecology and behaviour of the black-footed cat Felis
nigripes in the Kimberley area, South Africa. Pp. 8- 16
in G. Olbricht and U. Schurer, eds. International studbook for the black-footed cat, 1992. Zoologischer
Garten Stadt Wuppertal, Wuppertal, Germany.
- . 1994. Diet and feeding behaviour of the black-footedcat (Felis nigripes Burchell, 1824) in the Kimberleyregion, South Africa. 2001. Garten N.F. 64(2): 83-96.
Slough, B.G. and Jessup, R.H. 1994. Case study on theecological sustainability of consumptiveand nonconsumptiveusesof wild species-fur-bearerharvesting in
the Yukon territory, Canada. Paper presented at the
IUCN/SSC SpecialistGroup on the SustainableUseof
Wild Speciesmeeting, 12-14Jan. 1994,BuenosAires.
- and Ward, R.M.P. 1990.Lynx harvest study:- 1988/89
progressreport. Yukon Dept. of RenewableResources,
Whitehorse,Canada.
Small, R.L. 1971. Interspecific competition among three
speciesof Carnivora on the Spider Ranch, YavapaiCounty, Arizona. M.S. thesis,Univ. Arizona, Tucson.
Smallwood, K.S. and Fitzhugh, E.L. 1993. A rigorous
technique for identifying individual mountain lions
Felis concolor by their tracks. Biological Conservation
65:5 l-59. (Reprinted with kind permission from
Elsevier Science,Kidlington OX5 lGB, U.K.)
Smimov, M.N., Sokolov, G.A. and A.N. Zyryanov. 1990.
The snow leopard (Uncia uncia, Schreber 1776) inSiberia. Intl. Ped. Book of SnowLeopards6: 9-15.
Smit, C.J. and Wijngaarden, A. van. 1976. Threatened
mammalsin Europe. Council of Europe, Strasbourg.
Smith, A. 1972. Breeding endangered species. New
Scientist 11 May: 333-335.
Smith, A.T., Formozov, N.A., Hoffmann, R.S., Zheng, C.L. and M.A. Erbajeva. 1990. The pikas. Pp. 14-60 in
J.A. Chapman and J.E.C. Flux, eds. 1990. Rabbits,
haresandpikas: statussurvey and conservationaction
pEan.IUCN, Gland, Switzerland and Cambridge,U.K.
Smith, J.L.D. 1978. Smithsonian Tiger EcologyI Project
report no. 13. Unpubl. report, SmithsonianInstitution,
Washington,D.C.
- . 1984. Dispersal, communication and conservation
strategies for the tiger (Panthera tigris) in Royal
Chitwan National Park, Nepal. Ph.D. thesis, Univ.
Minnesota, St. Paul.
- and McDougal, C. 1991. The contribution of variancein lifetime reproduction to effective population size intigers. Conserv.Biol. 5(4): 484-490.
- 7Wemmer, C., and H.R. Mishra. 1987a. A tiger geographic information system: the first step in a global
conservation strategy. Pp. 464-474 in R.L. Tilson and
U.S. Seal,eds.Tigersof the world: the biology, biopolitics, management,and conservationof an endangered

Acta Theriol. Sinica 3: 181-187 (in Chinese).
Shibnev, Y. 1989. The deerhunter. BBC Wildlz$e 7: 528534.
Shieff, A. and Baker, J. 1987. Marketing and international
fur markets. Pp. 862-877 in M. Novak, J. Baker, M.
Obbard and B. Malloch, eds. Wild furbearer management and conservation in North America. Ministry of
Natural Resources, Toronto.
Shoemaker, AS. 199 1. International studbookfor rareleopards. Riverbanks Zoological Garden, Riverbanks,
South Carolina.
- . 1993. The status of the leopard, Panthera pardus, innature: a country by country analysis. Riverbanks
Zoological Garden, Columbia, South Carolina.
Shortridge, G. 1934. The mammals of South West Africa,
Vol. 1. William Heinemann, London.
Sieve& P.R. and Keith, L.B. 1985. Survival of snowshoe
hares at a geographic range boundary. J. Wildl.
Manage. 49: 854-866.
Silveira, L. In submission. Notes on the distribution andnatural history of the pampas cat, Felis coZocoZo, in
Brazil. Mammalia.
Simberloff, D.A. and Cox, J. 1987. Consequences and costsof conservation corridors. Conserv. Biol. l(1): 63-7 1.
-, and D.W. Mehlman. 1992. Movement corridors: conservation bargains or poor investments? Conserv. Biol.
6(4): 493-504.
Sinclair, A. and Smith, J. 1984. Do plant secondary compounds determine feeding preferences of snowshoe
hare? Oecologia 6 1: 403-4 10.
Singh, A. I98 1. Tara: a tigress. Quartet, London.
- . 1984. Tiger! Tiger! Jonathan Cape, London.
Singh, R.L. 1990. Results of tiger census in India. Cat
News 13:3. Bougy-Villars, Switzerland.
Sinha, S.P. 1987. Ecology of wildlife with special reference to the lion (Panthera leo persica) in Gir Wildlife
Sanctuary,
Saurashtra,
Gujurat.
Ph.D. thesis,
Saurashtra University, Rajkot.
Sitton, L.W. 1977. California mountain lion investigations
with recommendations for management. Unpubl.
report, California Dept. Fish and Game, Sacramento.
-, Sitton, S. and D. Weaver. 1978. Mountain lion predation on livestock in California. Cal-Neva 1978: 174- 186.
Skead, C.J. 1980. Historical mammal incidence in theCape Province. Vol 1: the western and northern Cape.
Dept. Nature and Envl. Conservation of the Provincial
Administration of the Cape of Good Hope, Cape Town.
Skinner, J.D. 1979. Feeding behaviour in caracal, Felis
caracal. J. Zool., Lond. 189: 523-525.
- and Smithers, R.H.N. 1990. The mammals of the southern African subregion, 2d edn. Univ. of Pretoria Press,
Pretoria.
Sladek, J. 1973. The quantitative composition of the foodof the wild cat Felis silvestris living in the West374

References

species. Noyes, Park Ridge, New Jersey.
- 7McDougal, C.W. and M.E. Sunquist. 1987b. Female
land tenure system in tigers. Pp. 97- 109 in R.L. Tilson
and U.S. Seal, eds. Tigers of the world: the biology,
biopolitics, management, and conservation of anendangered species. Noyes, Park Ridge, New Jersey.
- 7 McDougal, C. and D. Miquelle. 1988. Scent marking
in free-ranging tigers, Panthera tigris. Anim. Behav. 36:
l-10.
Smith, N. 1976. Spotted cats and the Amazon skin trade.
Oryx 17(4): 362-370.
Smith, R.M. 1977. Movement patterns and feeding behaviour of the leopard in the Rhodes Matopos National
Park, Rhodesia. ArnoZdia 8( 13): l- 16.
Smithers, R.H.N. 1975. Family Felidae. Part 8.1. pp. l-10
in J. Meester and H.W. Setzer, eds. The mammals ofAfrica:an identification
manual. Smithsonian
Institution, Washington, D.C.
- . 197 1. The mammals of Botswana. Mus. mem. Natl.
Mus. Monum. Rhod. 4.
- . 1978. The serval Felis serval Schreber, 1776. S. Afr.
J. Wildl. Res. 8: 29-37.
- . 1983. The mammals of the southern Africa subregion,
1st edn. Univ. of Pretoria Press, Pretoria.
- . 1986. South African red data book-terrestrial
mammals. South African National Scientific Programmes
Report No. 125, Council for Scientific and Industrial
Research, Pretoria.
- and V.J. Wilson. 1979. Check list and atlas of the
mammals of Zimbabwe Rhodesia. Mus. mem. Natl.
Mus. Monum. Rhod. 9: 1- 147.
Smuts, G.L. 1976. Population characteristics and recent
history of lions in two parts of the Kruger National
Park. Koedoe 19: 153-164.
- . 1978a. Effects of population reduction on the travels
and reproduction of lions in Kruger National Park.
Carnivore I : 6 l-72.
- 1978b. More sex ratio data on lions. Carnivore l(2): 1.
- * 1982. Lion. Macmillan, Johannesburg.
-, Whyte, I.J. and T.W. Dearlove. 1977. A mass capture
technique for lions. E. Afr. Wildl. J. 15: 81-87.
- 7 Hanks, J. and I.J. Whyte. 1978. Reproduction andsocial organization of lions from the Kruger National
Park. Carnivore l(1): 17-28.
Sokal, R.R. and Crovello, T.J. 1970. Am. Nat. 104: 127.
Sokov, A.I. 1990. The present status of the snow leopard(Uncia uncia) population in the SW Pamir-Altai
Mountains (Tajikistan).
Intl. Ped. Book of Snow
Leopards 6: 33-36.
Sommerlatte, M., Festetics, A., Von Berg, F.-Ch. 1980.
[Survey of lynx by tracking following reintroductionin Austria.] Pp. 3 18-337 in A. Festetics, ed. [The lynx inEurope.] Kilda, Greven(in German).
Sopin, L.V. 1977. [The snow leopard in Altai.] Pp. 143-

144 in V.E. Sokolov, ed. [Proc. Congr. All-Union
Therio. Sot.] 2: Nauka, Moscow (in Russian).
Soul& M.E. 1987a.Introduction. Pp. 1-10 in M.E. Soul&
ed. Viable populations,for conservation. Cambridge
Univ. Press,Cambridge.
- . 1987b. Where do we go from here? Pp. 175-184 in
M.E. Soul& ed. Viable populationsfor conservation.
CambridgeUniv. Press,Cambridge.
-.,ed. 1987~. Viable populations for conservation.
CambridgeUniv. Press,Cambridge.
- . 1989.Theory andstrategy.Pp. 91-104in W.E. Hudson,
ed. Landscapelinkages and biodiversity. Island Press
andDefendersof Wildlife, Washington,DC.
- andSimberloff, D. 1986.What do geneticsandecology
tell us about the design of nature reserves? Biol.
Conserv. 35: 19-40.
- 7Gilpin, M., Conway, W. and T. Foose. 1986.The millenium ark: how long a voyage, how many staterooms,
how many passengers?
Zoo Biol. 5: 101- 113.
Spencer,J.A. and Morkel, P. 1992.Serological survey ofserafrom lions in EtoshaNational Park. S.Afr. J. Wildl.
Rex- 7 van Dijk, A.A., Horzinek, M.C., Egberink, H.F.,
Bengis,R.G., Keet, D.F., Lindeque,M., Morkel, P. and
D.H.L. Bishop. In press.Incidenceof feline immunodeficiency virus reactive antibodiesin free-ranginglionsof the Kruger and Etosha National Parks in southern
Africa. OnderstepoortJ. Vet. Res.
Stahl, P. 1986. [The Europeanforest wildcat (Felis silvestrisSchreber, 1777): resourceexploitation and spatial organization.] Ph.D. thesis,Univ. Nancy, Nancy (inFrench).
- . 1993. [Statusof the wildcat (Fehs silvestris)in western
Europe.] Pp. 16-25in Proc. seminaron the biologyd andconservation of the wildcat (Felis silvestris), Nancy,
France, 23-25 September 1992. Council of Europe,
Strasbourg(in French).
- and Artois, M. 1991. Status and conservation of thewildcat in Europe and around the Mediterranean rim.
Council of Europe, Strasbourg.
- and Leger, F. 1992. [The European wildcat (Felis silvestris Schreber, I777).] In [Encyclopedia qf Frenchcarnivores.] SocieteFrancaisepour l’etude et la protection desMammiferes, Puceul (in French).
- 9Artois, M. and M.F.A. Aubert. 1988. [Spatial organization and displacementof adult forest wildcats (FeZis
silvestris Schreber, 1777)in Lorraine.] Terre et Vie 43:
113-132(in French).
Stander,P. 1990a.Noteson foraging habits of cheetah.S.
Afr. J. Wildl. Res.20(4): 130-132.
- . 1990b. A suggestedmanagementstrategy for stockraiding lions in Namibia. S.Afr. J. Wildl. Res.20(2):3743.
- . 1991. Demography of lions in the Etosha Nationall375

References

Bougy-Villars, Switzerland.
- and Stuart, T. 1989. Leopard in the lower Orange Riverbasin-a survey of their conservation status. Unpubl.
report, African Carnivore Survey, Nieuwoudtville,
South Africa.
- . 1991. Regional size variation and sexual dimorphismof the leopard. Cat News 15: 9. Bougy-Villars,
Switzerland.
- . 1992a. Southern, Central and East African mammals:a photographic guide. Struik, Cape Town.
- . 1992b. Are leopards choosey when scratching trees?
Cat News 16: 8. Bougy-Villars, Switzerland.
Stuart, C.T. and Wilson, V.J. 1988. The cats of southern
Africa. Chipangali Wildlife Trust, Bulawayo.
Stuart, C.T., Macdonald, I.A.W. and M.G.L. Mills. 1985.
History, current status and conservation of large mammalian predators in Cape Province, Republic of South
Africa. Biol. Conserv. 3 1: 7- 19.
Stubbe, M. and Chotolchu, N. 1968. [Mammals ofMongolia.] Mitt. Zool., Mus. Berlin 44: 5- 12 1 (inGerman).
Suminski, H.R. 1982. Mountain lion predation on domestic
livestock in Nevada. Proc. Vert. Pest Conf 10: 62-66.
Suminski, P. 1962. [Characters of the pure form of thewildcat Felis silvestris Schreber.] Arch.Sci. 15: 277-296(in French).
- . 1977. [On the problems of distinguishing between thewildcat, Felis silvestris Schreber, 1777, and the domestic cat, Felis catus Linnaeus, 1758.1 Saugetierk. Mitt.
40: 236-238 (in German).
Sunquist, F. 1991. The living cats. Pp. 28-53 in J.
Seidensticker and S. Lumpkin, eds. Great Cats.
Merehurst, London.
Sunquist, M.E. 198 1. The social organization of tigers(Panthera tigris) in Royal Chitwan National Park,
Nepal. Smithson. Contrib. 2001. 336: l-98.
- . 1983. Dispersal of three radiotagged leopards. J.
Mamm. 64: 337-341.
- . 1992. The ecology of the ocelot: the importance ofincorporating life history traits into conservation plans.
Pp. 117- 128 in Anon. 1992~. [Felids of VeneZuela.1[Foundation for the Study of Physical, Mathematicaland Natural Sciences (FUDECI)], Caracas.
- and Sunquist, F.C. 1989. Ecological constraints on predation by large felids. Pp. 283-301 in J.L. Gittleman,ed. Carnivore behavior, ecology and evolution.
Chapman and Hall, London.
- 7 and D.E. Daneke. 1989. Ecological separation in a
Venezuelan llanos carnivore community. Pp. 197-232
in K.H. Redford and J.F. Eisenberg, eds. Advances in
Neotropicalmammalogy. Sandhill Crane Press,
Gainesville, Florida.
*
Sunquist, M., Leh, C., Sunquist, F., Hills, D.M. and R.
Rajaratnam. 1994a. The Bomean bay cat. Cat News 20:

Park. Madoqua 18: l-9.
- . 1992a. Foraging dynamics of lions in a semi-arid environment. Can. J. 2002. 70.
- . 1992b. Cooperative hunting in lions: the role of theindividual. Behav. Ecol. Sociobiol. 29: 445454.
- . 1993. Conservation of African carnivores in a developing world. In W. van Houven and H. Ebedes, eds.
Wildlife ranching: a celebration ofdiversity. Proc. 3rd
Int. Wildl. Ranching Symp.,
Pretoria, October 1992.
w
National Game Organization, Pretoria.
- and Albon, S.D. 1992. Hunting success of lions in asemi-arid environment. In N. Dunstone and M.L.
Gorman, eds. Mammals as predators. Proc. Symp.
2001. Sot. Land. 65. Clarendon, Oxford.
- and Stander, H. 1987. Characteristics of lion roars in
Etosha National Park. Madoqua 15: 3 15-3 18.
Stehlik, J. 197 1. Breeding jaguars Panthera onca at
Ostrawa Zoo. Int. Zoo Yearb. 11: 116- 118.
Stephenson, R.O. 1986. Development of lynx population
estimation
techniques. Unpubl. report, Alaska
Department of Fish and Game, Juneau, Alaska.
-, Grangaard, D.V. and J. Burch. 199 1. Lynx, Felis lynx,
predation on red foxes, Vulpes vulpes, caribou,
Rangifer tarandus, and Dal1 sheep, Ovis dalli, in
Alaska. Can. Field-Nat. 105: 255-262.
Sterndale, R.A. 1884. Natural history of the mammalia ofIndia and Ceylon. Himalayan Books, New Delhi (1982
reprint).
Stevens, WK. 1994. Threat of encroaching deserts may be
more myth than fact. New York Times 18 Jan: B5.
Stockley, C.H. 1936. Stalking in the Himalayas and northern India. London.
Storer, T.I. 1923. Rabies in a mountain lion. California
Fish and Game 9(2): 45-48.
Stroganov, S.U. 1962. Carnivorous mammals of Siberia.
Engl. transl., Israel Program for Scientific Translation,
Jerusalem, 1969.
Stuart, C.T. 1977. Analysis of Felis lybica and Genetta
genetta scats from the Central Namib Desert, SouthWest Africa. Zoologica Africana 12: 239-241.
- . 198 1. Notes on the mammalian carnivores of the Cape
Province, South Africa. Bontebok 1: l-58.
- . 1982. Aspects of the biology of the caracal (Felis caracal) in the Cape Province, South Africa. M.S. thesis,
Univ. Natal, Pietermaritzburg.
- . 1984. The distribution and status of Felis caracal
Schreber, 1776. Siiugetierk. Mitt. 3 1: 197-203.
- . 1985. The status of two endangered carnivores occurring in the Cape Province, South Africa, Felis servaland Lutra maculicollis. Biol. Consew. 32: 375-382.
- . 1986. The incidence of surplus killing by Panthera
pardus and Felis caracal in Cape province, South
Africa. Mammalia 50(4): 556-558.
- . 199 1. Lion. Unpubl. data sheet, Cat Specialist Group,376

References13- 15. Bougy-Villars, Switzerland.
- . 1994b. Rediscovery of the Bornean bay cat. Oryx28( I): 67-70.
Suzuki, H., Hosoda, T., Sakurai, S., Tsuchiya, K.,
Munechika, I., and Korablev, V. 1994. Phylogenetic
relationship between the Iriomote cat and the leopard
cat, FeZis Den&ens&
based on the ribosomal DNA.
Jpn. J. Genet. 69:397-406.
Swank, W.G. and Teer, J.G. 1987. Status ofthe jaguar.
Unpubl. report, National Fish and Wildlife Foundation,
Washington, D.C.
- . 1988. A proposed jaguar country management plan.
Unpubl. report, Safari Club International, Reston,
Virginia.
Sweanor, L.L. 1990. Mountain lion social organization ina desert environment. M.S. thesis, Univ. of Idaho,
Moscow.
- and Logan, K.A. 1992. Life among desert cougars. New
Mexico Wildl. Nov-Dee: 2-26.
Sweeney, S.J. 1978. Diet, reproduction and population
structure qf the bobcat (Lynx rufus fasciatus) in western
Washington. M.S. thesis, Univ. Washington, Seattle.
Swift, J. 1975. The Sahara. Time-Life Books, Amsterdam.
Swinhoe, R. 1862. The mammals of Formosa (Felidae).
Proc. 2001. Sot. Lond. 1862: 352-353.
Szemethy, L. 1989. Occurrences and densities of wild cat
and badger in Hungary. Vaobiologia 3: 163- 168.
- . 1993. The actual status of wildcat (Felis silvestris) in
Hungary. Pp. 52 in Proc. seminar on the biology” andconservation of the wildcat (Felis silvestris), Nancy,
France, 23-25 September 1992. Council of Europe,
Strasbourg.
Taber, R., Miller, S. and J. Rottmann. 1974. Rare andendangered mammals of Chile. Unpubl. report, World
Wildlife Fund, Morges.
Tan Bangjie. 1984. The status of felids in China. Pp. 33-47
in P. Jackson, ed. The plight of the cats: proc. meeting
and workshop of the KJCN/SSC Cat Specialist Groupat Kanha National Park, Madhya Pradesh, India, 9-12
April 1984. Unpubl. report, IUCN/SSC Cat Specialist
Group, Bougy-Villars, Switzerland.
- . 1987. Current status of Chinese tigers. Cat News 6: 710. Bougy-Villars, Switzerland.
Tatarinov, K.A. 1983. [Mammals of Kabardino-Balkaria.][Proc. Congr. All-Union Therio. Sot.] 3: 182- 183.
Nauka, Moscow (in Russian).
Taylor, C.R. and Rowntree, V.J. 1973. Temperature regulation and heat balance in running cheetahs: a strategy
for sprinters. Amer. J. Physiol. 224: 848-851.
-, Shkolnik, A., Dmtel, R., Baharav, D. and A. Borut.
1974. Running in cheetahs, gazelles, and goats: energy
cost and limb configuration. Am. J. Physiol. 227(4):848-850.
Teer, J.B. and Swank, W.G. 1977. Status of the leopard

in Africa south ofthe Sahara. Unpubl. report, U.S. Fish
and Wildlife Service, Washington, D.C.
Tehsin, R. and F. Tehsin. 1990. Jungle cat Fehs chaus andgrey junglefowl Gallus soonerati. J. Bombay.I Nat. Hist.
Sot. 87: 144.
Tello, J. 1986a. Survey ofprotected areas and wildlife
areas in Mozambique, with recommendations for theirconservation. Unpubl. report, WWF International,
Gland, Switzerland.
Tello, J.L. 1986b. The situation ofthe wild cats (Felidae) in
Bolivia. CITES Secretariat, Lausanne.
Templeton, A. 1989. Pp. 3-27 in D. Otte and J.A. Endler,
eds. Speciation and its consequences.
Sinauer,
Sunderland, Massachusetts.
Terborgh, J. 1988. The big things that run the world-asequel to E.O. Wilson. Conserv. Biol. 2(4): 402-403.
Tewes, M.E. 1986. Ecological and behavioral correlates
ofocelot spatial patterns. Ph.D. thesis, Univ. Idaho,
Moscow.
- and Everett, D.D. 1986. Status and distribution of theendangered ocelot and jaguarundi in Texas. Pages 147158 in S.D. Miller and D.D. Everett, eds. Cats oftheworld:
biology, conservationand management.
National Wildlife Federation, National Wildlife
Federation.
- and Scott, J.M. 1987. Assessment ofbobcat harvest in
the western
United States, 1975-l 985. Idaho
Cooperative Wildlife Research Unit, Univ. of Idaho,
Moscow.
Thapar, V. 1986. Tiger: portrait of a predator. Collins,
London.
- . 1989. Tigers: the secret life. Elm Tree Books, London.
- . 1992. The tiger’s destiny. Kyle Cathie, London.
Thomas, C.D. 1990. What do real population dynamics tell
us about minimum viable population sizes? Conserv.
Biol. 4(3): 324-327.
Thomas, E.M. 1990. The old way. New Yorker Ott 15: 78110.
- . 1994. The tribe oftiger: cats and their culture. Simonand Schuster, New York.
Thomsen, J. and Luxmoore, R. 1990. Sustainable utilization of wiZdZi$efor trade. Paper presented to the IUCN
18th General Assembly, Perth, Australia.
Thornback, J. and Jenkins, M., eds. 1982. ZUCN mammal
red data book, part I. IUCN, Gland, Switzerland andCambridge, U.K.
Thorne, E.T. and William, E.S. 1988. Conserv. Biol. 2(l):66-74.
Thresher, P. 1982. The lion’s share. New Scientist15 Apr.
Tikader, B.K. 1983. Threatened animals of India.
Zoological Survey of India, Calcutta.
Tiger Trust. 1993. Bones of contentionpt. 2: the tiger trade
continues. Unpubl. report, Tiger Trust, Bury St.
Edmunds, U.K.

377

References

Town.
Turyanin, 1.1.1988. [Felids at the Ukrainian Carpathians.]
Pp. 91-95 in [Investigations of the Ukrainian theriofauna and its managementand conservation.] Kiev (inRussian).
Udvardy, M.D.F. 1975. A classification of the biogeographical provinces of. the world. IUCN Occasional
Paperno. 18, Morges.
Ullrich, B. and Riffel, M. 1993. New evidence for theoccurrenceof the Anatolian leopard,Panthera pardus
tulliana (Valenciennes, 1856), in western Turkey.
Zoology in the Middle East 8: 5- 14.
Ulmer, F.A., Jr. 1941.Melanism in the Felidae, with special reference to the genusLynx. J. Mammal. 22: 285288.
- . 1968. Breeding fishing cats, Felis viverrina, at
PhiladelphiaZoo. Int. Zoo. Yearb. 8: 49-55.
United StatesFish and Wildlife Service (USFWS). 1982.
Threatened statusfor the leopard in southern Africa.
Federal Register47(19): 4204-4211.
- . 1987.Florida Panther (Felis concolor coryi) Recoveryw
Plan. Prepared by the Florida Panther Interagency
Committee for the U.S. Fish and Wildlife Service,
Atlanta, Georgia.
- . 1993. Notice of intent to prepare an environmental
assessment(EA) addressingpotential genetic management options for the Florida panther. Federal Register58(157): 43651.
Ustay, A.H. 1990.Hunting in Turkey. BBA, Istanbul.
van Aarde, R.J. and van Dyk, A. 1986. Inheritance of theking coat colour patterns in cheetahsAcinonyx jubatus. J. 2001.209: 573-578.
- and Skinner, R.D. 1986. Pattern of spaceuseby relocated servalsFelis serval. Afr. J. Ecol. 24: 97- 101.
Van Bree, P.J.H. and Khan, M.K. 1992. On a fishing cat,
Felis (Prionailurus) viverrina Bennett, 1833,from continental Malaysia. 2. Saugetierk.57: 179-180.
van den Berg, M. and Damhuis,R. 1982.The fur trade in
Kashmir. TRAFFIC Bulletin: 59.
van den Brink, F.H. 1971. [The pardel lynx in France.]
Bull. Sot. d’Etude Sci. Nat. Nfmes 5 1: 109-117 (inFrench).
van Dyk, A. 1991. The cheetahsof DeWildt. Struik, Cape
Town.
van Dyke, F.G. 1983.A westernstudy of cougar track surveysand environmentaldisturbancesaflecting cougars
related to the status of the eastern cougar Felis concolor cougar. Ph.D. thesis, State Univ. New York,
Syracuse.
-, Brocke, R.H., Shaw, H.G., Ackerman, B.B., Hemker,
T.P. and F.G. Lindzey. 1986a.Reactionsof mountainlions to logging and humanactivity. J. Wildl. Manage.
50(l): 95-102.
-, Brocke, R.H. andH.G. Shaw. 1986b.Useof road track

Tilson, R. 1992a. A summary report on the Sunzatran tiger
population and habitat viability analysis workshop.
Unpubl. report, IUCN/SSC Captive Breeding Specialist
Group, Apple Valley, Minnesota.
- . 1992b. Cats in zoos. Pp. 214-219 in J. Seidensticker
and J. Lumpkin, eds. Great cats. Merehurst, London.
- 7Princee, F. and T.J. Foose. 1992. Tiger global animal
survival plan. Unpubl. draft circulated for review,
September 1992. Minnesota Zoological Gardens,
Apple Valley.
Tinsley, J.B. 1970. The Florida panther. Great Outdoors,
St. Petersburg, Florida.
- . 1987. Puma: legendary lion of. the Americas. Univ. of
Texas WesternPress,El Paso.
Tischendorf, J., ed. 1991. Cougar populations: their status, managementandfuture. Unpubl. report, Amer.
Ecol. Res.Inst., Fort Collins, Colorado.
- 7ed. 1992a.Easternpanther update1. Amer. Ecol. Res.
Inst., Fort Collins, Colorado.
- . 1992b. The easternpanther and the burden of proof.
Cat News 16: 13-14. Bougy-Villars, Switzerland.
- and Henderson, F.R. 1993. The puma in the central
mountains and plains. Paper presentedat the Wildl.
Sot. meeting, 4-5 Aug 1993,Manhattan, Kansas.
Todd, A.W. 1985.The Canadalynx: ecology andmanagement. CanadianTrapper 13(2): 15-20.
Todd, N.B. 1977.Cats and commerce.Sci. Am. 237: lOO107.
Tomkies, M. 1991. Wildcats. Whittet, London.
Tonkin, B. and Kohler, E. 1978. Breeding the African
goldencat in captivity. Int. Zoo Yearb. 18: 147-150.
Toweill, D.E. 1982.Winter foods of easternOregon bobcats.Northwest Sci. 56: 3 1O-315.
Toynbee, J.M.C. 1973. Animals in Roman life and art.
Thamesand Hudson,London.
Trainer, C.E. 1975.Direct causesof mortality in mule deerfawns duing summer and winter periods on Steens
Mountain, Oregon. Proc. West.Assoc. State Gameand
Fish Comm.55: 163-170.
Trinh Viet Cuong. 1991. [Somedata on the Carnivora in
the Tay Nguyen region.] M.S. thesis, Univ. Hanoi,
Hanoi (in Vietnamese).
Trulio, L. 1989.Livestock depredation:why lions are nota major threat to California’s livestock industry. In
Pouncing on the myths about mountain lions 4,
Mountain Lion Preservation Foundation Series,
Sacramento.
Tudge, C. 1976. Last animalsat the zoo. New Scientist 15
Jul: 134-136.
Tumlison, R. 1987. Felis lynx. Mammalian Species269:
1-8.
Tun Yin, U. 1967. Wild animals of Burma. Rangoon
Gazette Ltd., Rangoon.
Turnbull-Kemp, 1967. The leopard. Timmins, Cape

378

References

counts as indices of mountain lion presence. J. WilcJZ.
Manage. 50( 1): 102- 109.
van Gils, H.A.M.J. 1993. Vegetation classification:areview for harmonization of maps. Paper presented at
the UNEP-HEM/WCMC/GCTE
Preparatory Workshop on Vegetation Classification,Charlotteville,
Virginia, 24-26 Jan.
van Gruisen, J. and Sinclair, T. 1992. Fur trade in
Kathmandu: implications for India. TRAFFIC India,
New Delhi.
van Helvoort, B.E., de Iongh, H.H. and P.J.H. Van Bree.
1985. A leopard skin and skull (Panthera pardus L.)from Kangean Island, Indonesia. 2. Saugetierk. 50:
182- 184.
van Ingen and van Ingen. 1948. Interesting shikar trophies:
hunting leopard (A. jubatus). J. Bombay Nat. Hist. Sot.
47(4): 7 18.
Van Mensch, P.J.A. and Van Bree, P.J.H. 1969. On the
African golden cat, Profehs aurata (Temminck, 1827).
Biologia Gabonica 5: 235-269.
van Meter, V.B. 1988. The Florida panther. Pamphlet,
Florida Power and Light Co., Miami.
van Orsdol, K.G. 1982. Ranges and food habits of lions in
Ruwenzori National Park, Uganda. Symp. 2002. Sot.
Lond. 49: 325-340.
- . 1984. Foraging behaviour and hunting success of lions
in Queen Elizabeth National Park, Uganda. Afr. J. Ecol.
22: 79-99.
- 7Hanby, J.P. and J.D. Bygott. 1985. Ecological correlates of lion social organization (Panthera leo). J. Zool.,
Lond. 206: 97-112.
van Peenen, D.P.D. 1969. Preliminary identification manual,for the mammals ofSouth Vietnam. Smithsonian,
Washington, D.C.
van Saceghem, R. 1942. [A new species of felid fromAfrica, Felis maka. ] Bull. Sot. bot. ~001. Congo 5 : 1316 (in French).
van Sickle, W.D. and Lindzey, F.G. 1992. Evaluation ofroad track surveys for cougars (Felis concolor). GreatBasin Nat. 52(3): 232-236.
van Zyll de Jong, C.G. 1963. The biology ofthe lynx, Felis(Lynx) canadensis Kerr in Alberta and the Mackenzie
District,
N. W.T. M.S. thesis, Univ. of Alberta,
Edmonton.
- . 1966. Food habits of the lynx in Alberta and theMackenzie District, N.W.T. Canadian Field-Nat. 80:
18-23.
- . 197 1. The status and management of the Canada lynx
in Canada. Pp. 15-22 in S.E. Jorgenson and L.D. Mech,eds. Proc. symp. on the native cats ofNorth America:
their status and management. U.S. Fish and Wildlife
Service, Twin Cities.
Vasiliu, G.D. and Decei, P. 1964. [On lynx in the
Romanian Carpathian Mountains.] Saugetierk. Mitt. 12:

155- 183 (in German).
Vaughan, C. 1983. A report on denseforest habitat forendangered wildlife species in Costa Rica. Unpubl.
report, Universidad National, Heredia.
- . In press. The mysterious jaguarundi. Anima (Japan).
Vereshchagin, N.K. 1959. The mammals of the Caucasus:
a history ofthe evolution of thefauna. Nauka, Moscow.
Engl. transl. by Israel Program for Scientific Translations, Jerusalem, 1967.
Verma, K. 1983. The skin haven. India Today Nov 15:110.
Vigne, J.D. 1992. Zooarchaeology and the biogeographical history of the mammals of Corsica and Sardinia
since the last Ice Age. Mammal Review 22: 87-96.
Viljoen, P.C. 1993. The effects of changes in prey availability on lion predation in a large natural ecosystem
in northern Botswana. In N. Dunstone and M.L.
Gorman, eds. Mammals as predators. Proc. Svmp.
2001. Sot. Lond. 65. Clarendon, Oxford.
Villafuerte, R. and Moreno, S. 199 1. Rabbit haemorragic
disease (RHD) in Donana National Park (DNP). Trans.
Int. Union Game BioEogists 20: 107-108.
Visser, J. 1977. The small cats. African WildlifeI 3 l(1):26-28.
- . 1978. Status and conservation of the smaller cats ofsouthern Africa. Pp. 60-66 in R.L. Eaton, ed. The
world’s cats 3(1): contributions to status, management
and conservation, 2d edn. Carnivore Res. Inst., Univ.
Washington, Seattle.
Vorster, F. 1988. Vraelysopname van die roofdierverwant
kleinveeverliese in die Nuwe-Roggeveld
Afdelingsraadgebied van die Kaapprovinsie. [Survey of predator-related small stock losses in the Nuwe-Roggeveld
district of the Cape province.] Bontebok 6: 26-29 (inAfrikaans with Engl. summary).
Vuosalo, E. 1976. Once there was a tiger. Wildlife Mar:
126- 130 (Teheran).
Walker, S. 1990. The king retreats: from his sub-continental hunting grounds the Asiatic lion has been pushed
into the restricted environs of the Gir. Zllustrated
Weekly of India, 2 September 1990.
- . 1994. Executive summary of the Asiatic lion PHVA.
First draft report. Zoo’s Print Jan/Feb: 2-22
(Coimbatore, India).
Wallach, J.D. and Boever, W.J. 1983. Diseases of. exotic
animals: medical and surgical management. W.B.
Saunders, Philadelphia.
Walton, B. 1991. Catcalls. BBC Wildlife 9(3): 198-202.
Wang Qishan, ed. 1990. [The mammals ofAnhui.1 Hefei(in Chinese).
Wang Zongyi and Wang Sung. 1986. Distribution andrecent status of the Felidae in China. Pp. 201-210 in
S.D. Miller and D.D. Everett, eds. Cats of. the world:
biology, conservation and management. National379

References

13: 18. Bougy-Villars, Switzerland.
West, S. 199 I. Preserving cougar country: a guide to protecting mountain lion and deer habitat in California.
Unpubl.
report,Mountain
Lion Foundation,
Sacramento.
Wharton, C.H. 1968. Man, fire and wild cattle in southeast Asia. Proc. Ann. Tall Timbers Fire Ecol. Con& 8:
107-167.
Wharton, D. and Freeman, H. 1988. The snow leopard in
North America: captive breeding under the Species
Survival Plan. Pp. 13 I- 136 in H. Freeman, ed. Proc.
fifth international snow leopard symposium. Wildlife
Institute of India and International Snow Leopard Trust,Seattle.
Whittle, R.K. 1979. Age in relation to the winterfood
habits and helminth parasites of the bobcat (Lynxrufus, Schreber) in Oklahoma. M.S. thesis, Oklahoma
State Univ., Stillwater.
Whyte, I.J. and Joubert, S.C.J. 1988. Blue wildebeest population trends in the Kruger National Park and theeffects of fencing. S. Afr. J. Wildl. Res. 18(3): 78-87.
Widholzer, F.L., Bergmann, M. and C. Zotz. 198 1.
Breeding the little spotted cat. Int. Zoo News 28: 17-22.
Wiese, R.J. and Hutchins, M. 1993. The role of captive
breeding and reintroduction in wildlife conservation. In
Endangered wildlife of the world. Marshall Cavendish,
New York.
Wildt, D.E. 1991a. Fertilization in cats. Pp. 299-328 in
B.S. Dunbar and M. O’Rand, eds. A comparative
overview of mammalian fertilization. Plenum, NewYork.
- . 199 1b. Biotechnology for animal conservation: potential and limitations. Pp. 87-92 in J.G. Grootenhuis, S.G.
Njuguna and P.W. Kat, eds. Wildlife research for sustainable development: proc. conf 22-26 April 1990.
Kenya Agricultural Research Institute, Kenya Wildlife
Service, and National Museums of Kenya, Nairobi.
- . 1991~. Advances in artificial reproduction. Pp. 220221 in J. Seidensticker and J. Lumpkin, eds. Great cats.
Merehurst, London.
- . 1992a. Genetic resource banking for conserving
wildlife species: justification, examples and becoming
organized on a global basis. Anim. Reprod. Sci 28: 247257.
- . 1992b. Reproductive strategies for understanding,managing and conserving wild felids. Unpubl. projectprogress report submitted to Felid Taxon Advisory
Group.
-, Schiewe, M.C., Schmidt, P.M., Goodrowe, K.L.,
Howard, J.G., Phillips, L.G., O’Brien, S.J. and M.
Bush. 1986. Developing animal model systems forembryo technologies in rare and endangered wildlife.
Theriogenology 25: 33-5 1.
-, O’Brien, S.J., Howard, J.G., Caro, T.M., Roelke, M.E.,

Wildlife Federation, Washington, D.C.
Ward, R.M.P. and Krebs, C.J. 1985. Behavioural
responses of lynx to declining snowshoe hare abundance. Can. J. 2001. 63: 2817-2824.
Watson, R.W.M. 1980. Golden cat in the Aberdare Forest.
East Afr. Nat. Hist. Sot. Bull. 1980: 14.
Wayne, R.K., Benveniste, R.E., Janczewski, D.N. and S.J.
O’Brien. 1989. Molecular and biochemical evolution of
the Carnivora. Pp. 465-494 in J.L. Gittleman, ed.
Carnivore behavior, ecology and evolution. Cornell
Univ. Press, New York.
-, Lehman, N. and S.M. Jenks. 1992. The use of morphologic and molecular techniques to estimate genetic
variability and relationships of small populations. Pp.
217-235 in D.R. McCullough and R.H. Barrett, eds.
Wildlife 2001: populations. Elsevier, New York andLondon.
Weber, C. 1983. [Representation of the biogeographicalprovinces in biosphere reserves of Chile. Present andfuture actions of the Corporation Forestal National.]
Boletl’n Tecnico 10. Corporation National Forestal,
Ministerio de Agricultura, Santiago (in Spanish).
Weigel, I. 196 1. Das Fellmuster der wildlebenden
Katzenarten und der Hauskatze in vergleichender
Stammesgeschichtlicher Hinsicht. [The pelage patternsof wild-living cat species and domestic cats comparedwith aspects of phylogenetic history.] Saugetierk. Mitt.
9(Suppl: l-120).
- . 1975. Small cats and clouded leopards. Pp. 281-332
in B. Grzimek, ed. Grzimek’s animal life encyclopedia
12: MammaZs III. Van Nostrand Reinhold, New York.
Weisbein, Y. 1989. [The biology and ecology of the caracal (Felis caracal) in the Aravah Valley of Israel.] M.S.
thesis, Tel Aviv Univ., Tel Aviv (in Hebrew withEnglish summary).
Weisbein, Y. and Mendelssohn, H. 1990. The biology andecology of the caracal Felis caracal in the northern
Aravah Valley of Israel. Cat News 12: 20-22. BougyVillars, Switzerland.
Weiss, G. 1952. [Observations on two separately raised
housecats.] Zschr. f Tierpsychol.
9: 45 l-462 (inw
German).
Wemmer, C. and Sunquist, M.E. 1988. Felid reintroductions: economic and energetic considerations. In H.
Freeman, ed. Proc. fifth international snow leopard
symposium. Wildlife Institute of India and International
Snow Leopard Trust, Seattle.
Werdelin, L. 198 1. The evolution of lynxes. Ann. 2001.
Fenn. 18: 37-71.
- . 1983a. Morphological patterns in the skulls of cats.
Biol. J. Linn. Sot. 19: 375-391.
- . 1983b. Small Pleistocene felines of North America. J.
Vertebrate Paleontology 5: 194-2 10.
- . 1990. Taxonomic status of the pardel lynx. Cat News

380

References

Woloszyn, D. and Woloszyn, B.W. 1982. [The mammalsof Sierra de La Laguna, Baja California Sur.] Consejo
National de Ciencia y Technologia, Mexico (inSpanish).
Won Hong-Koo. 1968. [Mammals of Korea.] Nat. Acad.
Sci, Pyongyang (in Korean).
Won Pyong-Oh. 198 1. [Red Data Book of the Republic ofKorea: rare and endangered species of animals andplants.] Korean Assn. for the Conservation of Nature(in Korean with Engl. summary).
- . 1988. Rare and endangered species of animals in South
Korea. Bull. Institute of Ornithology, Kyung Hee Univ.
2: 61-65.
Woodford, M.H. and Rossiter, P.B. 1993. Diseaserisks
associatedwith wildlife translocation projects. Rev.
sci. tech. 08 int. Epiz. 12(l): 115-135.
Worah, S. 1991. The ecology and managementof a fragmentedforestin southGujarat, India: the Dangs.Ph.D.
thesis,Univ. of Pune, Pune.
World Conservation Monitoring Centre (WCMC). 1992.
Global biodiversity: status of the Earth’s living
resources.B. Groombridge,ed. Chapmanand Hall and
IUCN, London.
World Resources Institute (WRI), United Nations
Environment Programme(UNEP), and United Nations
Development Programme (UNDP). 1992. World
Resources1992-1993.Oxford Univ. Press,New York.
Wozencraft, W.C. 1993.Order Carnivora. Pp. 286-346 in
D.E. Wilson and D.M. Reeder, eds.Mammal speciesof the world, 2d edn. Smithsonian,Washington,D.C.
Wright, A. 1984.A note on the wild cats of the northeastern region of India. Pp. 81-84 in P. Jackson, ed. The
plight of the cats: proc. meeting and workshop of the
IUCN/SSC Cat Specialist Group at Kanha National
Park, Madhya Pradesh, India, 9-12 April 1984.
Unpubl. report, IUCN/SSC Cat Specialist Group,
Bougy-Villars, Switzerland.
Wright, B. 1989.A glimpseof tiger family life. Cat News
11: 16. Bougy-Villars, Switzerland.
Wright, B.S. 1972. The easternpanther-a question ofsurvival. Clarke Irwin, Toronto.
- . 1959. The ghost of North America-the story* of theeasternpanther. Vantage, New York.
Wright, S. 1969. Evolution and the genetics ofpopula.
tions,vol. 2: Theory of genefrequencies.Univ. Chicago
Press,Chicago.
Wroe, D.M. and Wroe, S. 1982. Observations of bobcat
predationon bats.J. Mammal. 63: 682-683.
Wrogemann, N. 1975. Cheetah under the sun. McGrawHill, Johannesburg.
Wurster-Hill, D.H. and Gary, C.W. 1975.The inter-relationship of chromosomebanding patterns in procyonids, viverrids, and felids. Cytogen. Cell Genet. 15:
306-331.

Brown, J.L. and M. Bush. 1987a. Similarity in ejaculate-endocrine characteristics in captive versus freeranging cheetahs of two subspecies. Biol. Reprod. 36:
35 l-360.
- 7Bush, M., Goodrowe, K.L., Packer, C., Pusey, A.E.,
Brown, J.L., Joslin, P. and S.J. O’Brien. 1987b.
Reproductive and genetic consequences of founding
isolated lion populations. Nature 329: 328-33 1.
- 7 Phillips, L.G., Simmons, L.G., Chakraborty, PK.,
Brown, J.L., Howard, J.G., Teare, A., and Bush, M.
1988. A comparative analysis of ejaculate and hormonal characteristics of the captive male cheetah, tiger,
leopard and puma. Biol. Reprod. 38:245-255.
-, Mellen, J.D. and U.S. Seal, eds. 1992a. Felid action
plan, 1991 and 1992: AAZPA Felid Taxon Advisory
Group regional collection plan and IUCN Captive
Breeding Specialist Group global felid action plan.
National Zoological Park, Front Royal, Virginia.
- 9Monfort, S.L., Donoghue, A.M., Johnston, L.A. andJ.G. Howard. 1992b. Embryogenesis in conservation
biology-or
how to make an endangered species
embryo. Theriogenology 37: 16 1- 184.
- 7Brown, J.L., Bush, M., Barone, M.A., Cooper, K.A.,
Grisham, J. and J.G. Howard. 1993a. Reproductive status of cheetahs (Acinonyxjubatus)
in North American
zoos: the benefits of physiological surveys for strategic planning. Zoo Biol. 12: 45-80.
- 7 Byers, A.P., Howard, J.G., Wiese, R., Willis, K.,
O’Brien, S.J., Block, J., Tilson, R.L. and W.F. Rall.
I993b. Tiger genome resource banking (GRB) action
plan: global need and a plan for the North American
region. Review and discussion draft, AAZPA, Tiger
SSP and CBSG, Apple Valley, Minnesota.
Wiley, E.O. 1990. Syst. 2001. 27: 17.
Williamson, D. and Williamson, J. 1985. Botswana’s
fences and the depletion of Kalahari wildlife. ParkslO(2): 5-7.
Willis, K. 1993. Use of animals with unknown ancestries
in scientifically managed breeding programmes. Zoo
Biol. 12: 161-172.
- and Wiese, R.J. 1994. Genetic variation maintenancestrategies for captive breeding programmes.
J.
Aquariculture and Aquatic Science (in press).
Winter-Blyth, M.A. 1949. The Gir forest and its lions. J.
Bombay Nat. Hist. Sot. 48: 494-514.
- and Dharmakumarsinhji, K.S. 1950. The Gir forest andits lions. J. Bombay Nat. Hist. Sot. 49: 456-470.
Wirth, R. 1990a. Reintroduction-sometimes
a conservation problem? Internat. Zoo News 22: 13- 17.
- . 1990b. Observations on Amur and “Amur” leopards.
Cat News 13: 10. Bougy-Villars, Switzerland.
Wolff, J. 1980. The role of habitat patchiness in the population dynamics of snowshoe hares. Ecol. Monogr. 50:
111-130.

381

References

York, W. 1973. A study of serval melanism in the
Aberdaresand somegeneral behavioral observations.
Pp. 191-197 in R.L. Eaton, ed. The world’s cats 1.
World Wildlife Safari, Winston, Oregon.
Young, S.P. 1958. The bobcat ofNorth America: its history, rife, habits, economicstatusand control. Wildlife
ManagementInstitute, Washington,D.C.
- and Goldman, E.A. 1946. The puma, mysterious
American cat. American Wildlife Institute, Washington, D.C.
Yu, J. and Wozencraft, W.C. In press.Prionailurus bengalensis. Mammalian Species. American Society of
Mammalogists.
Yukhi, N. and O’Brien, S.J. 1990. DNA variation of the
mammalianmajor histocompatibility complex reflects
genomic diversity and population history. Proc. Nat.
Acad. Sci. USA 87: 836-840.
Yunker, C.E. and Guirgis, S.S. 1969. Studies of rodent
burrows and their ectoparasitesin the Egyptian desert.
1. Environment and microenvironment: somefactors
influencing acarine distribution. J. Egypt. Publ. Hlth.
Assn. 44: 498-542.
Zeuner, F.E. 1963. A history of domesticated animals.
Harper andRow, New York.
Zezulak, D.S. and Schwab, R.G. 1979. A comparisonof
density, home range and habitat utilization of bobcat
populations at Lava Beds and Joshua Tree National
Monuments, California. Bobcat Res. Con5Natl. Wildl.
Fed. Sci. Ser. 6: 74-79.
Zhang, Y.-Z. 1991. Mammalian zoogeography and conservation of the endangeredspeciesin arid areasof
China. Pp. 269-284 in J.A. McNeely and V.M.
Neronov, eds.Mammalsin the Palearctic desert:status
and trendsin the Sahara-Gobianregion. RussianAcad.
Sci and Russian Committee for the UNESCO programmeon Man and the Biosphere(MAB), Moscow.
Zheltuchin, A.S. 1984. [Daily activity and sizesof homerangesof the European lynx in the southern taiga of
the upper Volga river region.] Bjull. Mosk. obsc. ispty.
prir., otd. biol. 89: 54-62 (in Russian).
- . 1992.Distribution and numbersof lynx in the Soviet
Union. Pp. 19-22in The situation, conservation needs
and reintroduction of lynx in Europe. Proc. symp. I719 October, Neuchatel. Council of Europe, Strasbourg.
Zhirjakov, V.A. 1990. On the ecology of the snow leopard (Uncia uncia) in the Zailisky-Alatau (Northern Tien
Shan).Intl. Ped. Book of SnowLeopards6: 25-30.
Zhong, W., Zhou, Q. and Sun, C. 1985. [The basic characteristics of the rodent pestson the pasture in Inner
Mongolia and ecological strategiesfor control.] Acta
Theriol. Sinica 5: 241-249(in Chinese).
Ziesler,G. 1992.Souvenir d’un chatdesAndes. [Encounter
with an Andean mountaincat.] Animan 50: 68-79.

-

and Centerwall, W.R. 1982. The interrelationships ofchromosome banding patterns in canids, mustelids,
hyena, and felids. Cytogenet. Cell Genet. 34: 17% 192.
-, Doi, T., Izawa, M. and Y. Ono. 1987. Banded chromosome study of the Iriomote cat. J. Heredityw 78: 105 107.
Ximenez, A. 1961. [New subspecies of pampas cat from
Uruguay, Felis colocolo muiioai.] Corn. 2001. Mus.
Hist. Nat. Montevideo 5(88): l-8 (in Spanish).
- . 1973. [Notes on neotropical Felidae III. Contributions
to the knowledge of the Geoffroy’s cat Felis geofroyi
D’Orbigny & Gervais, 1844 and its geographic forms(Mammalia, Felidae).] Papeis Avulsos de Zoologia,
Sao Paolo 27(3): 3 l-43 (in Spanish with Engl. summa@.
~ . 1975. Felis geofroyi. Mammal. Species 54: l-4.
- . 1988. [Notes on neotropical felids IX: Felis(Leopardus) pardalis mitts.] Comm. 2001. Mus. Hist.
Nat. Montevideo 12( 168): l-7 (in Spanish).
Yalden, D.W. 1993. The problems of reintroducing carnivores. In N. Dunstone and M.L. Gorman, eds.
Mammals as predators. Proc. Svmp. Zool. Sot. London
65. Clarendon, Oxford.
I- 7 Largen, M.J. and D. Kock. 1980. Catalogue of the
mammals of Ethiopia. 4. Carnivora. Ztalian J. Zool. 8:
169-272.
Yamada, J.K. and Durrant, B.S. 1989. Reproductive parameters of clouded leopards (Neofelis nebulosa). Zoo
Biol. 8: 223-23 1.
Yamaya, S. and Yasuma, S. 1986. Coleoptera from droppings of the Iriomote cat, Prionailurus iriomotensis(Imaizumi). Pp. 18 I- 193 in Papers on entomologypresented to Proj1 Takehiko Nakane in commemoration of
his retirement. Japan Society of Coleopterology,
Tokyo.
Yanez, J.L., Cardenas, J.C., Gezelle, P. and F.M. Jaksik.
1986. Food habits of the southernmost mountain lions(FeZis concolor) in South America: natural versus livestocked ranges. J. Mamm. 67: 604-606.
Yasuma, S. 198 1. Feeding behaviour of the Iriomote cat(Prionailurus
iriomotensis Imaizumi, 1967). Bull.
Tokyo Univ. Forests 70: 8 I- 140 (in Japanese).
- . 1984. The Iriomote cat. Pp. 49-54 in P. Jackson, ed.
The plight of. the cats: proc. meetingand workshop of
the IUCN/SSC Cat SpecialistGroup at Kanha National
Park, Madhya
I Pradesh, India, 9- 12 April 1984.
Unpubl. report, IUCN/SSC Cat Specialist Group,
Bougy-Villars, Switzerland.
- . 1988.Iriomote cat: king of the night. Animal Kingdom91(6): 12-21.
- and Alikodra, H.S. 1990.Mammals of Bukit Soeharto
Protection Forest. Tropical Rain Forest ResearchCenter (PUSREHUT), Universitas Mulawarman,
Samarinda,Kalimantan, Indonesia.

382

IUCN/SSC

ActionPlans for the Conservationof BiologicalAc*tiou Plm .fijr Af?icwil Pril7mtc Co~7sul-\‘crtiori: I Y&6- I YYO. Compiled
by J.F. Oates and the IUCN/SSC
Primate Specialist Group, 1986.
41 pp. (Out of print.)
Action Plcul jilr A.vim Primate Corl,sc-‘r-\,lltioil: I Y&7- 1 YYI. Compiled
by A.A. Eudey and the IUCN/SSC
Primate Specialist Group. 1987,
65 pp. (Out of print.)
Alltclopvs.
Glohul Sunq~ lrlrd Rqioml
Action Plasm. Part I. East
md Northeust A.fricu. Compiled by R. East and the IUCN/SSC
Antelope Specialist Group, 1988, 96 pp. (Out of print.)
Dolphir~s. Porpoisc~s mid Wh~11c.s. A11 Actiorr Plllll .f?jr- the Coll.Yt’Yl’CAtiOllof‘Biologicw1 IX\~ersit\~: 1 Y88- 1YY.Z Second Edition. Compiled by
W.F. Perrin and the IuCN/SSC
Cetacean Specialist Group. 1989,
27 pp. (Out of print.)
The Kouprq.
AI1 Ac.tiorl Plm ,for its Coll.sc-‘r1’I1tiotl. Compiled by J.R.
MacKinnon,
S.N. Stuart and the IUCN/SSC Asian Wild Cattle
Specialist Group, 1988, 19 pp. (Out of print.)
Wkisels, Ci\‘t>ts, Mori,~oo.sus cirzd their Relcrti\,c).s. A/i Ac.tiori Plm,for the Comenutiorz
of’kfustelids md Vi\*c>rrid.s. Compiled
by A. Schreiber, R. Wirth, M. Riffel, H. van Rompaey and the
IUCN/SSC Mustelid and Viverrid Specialist Group. 1989, 99 pp.
(Out of Print.)
A~lte1oIw.s. Globctl Sur\q~ culd Regioul Actiorz Plums. Put-t 2.
Southem md South-c.ul~trcll Afiiur. Compiled by R. East and the
IUCN/SSC Antelope Specialist Group. 1989, 96 pp. (Out of print.)
Asia

Rhimx
A/l Ac-tim Plm ,filr-their CorlsL’1.~~trtioII. Compiled by
Mohd. Khan bin Momin Khan and the IUCN/SSC Asian Rhino
Specialist Group, 1989, 23 pp. (Out of print.)

Tortoise.s umi Fresh~uterTurtles. AIM Ac’tioll Plm ,fbr theirCorls~~r7’lltic~r?. Compiled by the IUCN/SSC Tortoise andFreshwater Turtle Specialist Group. 1989, 47 pp.
~fj-iula Elephtrrlts mld Rhit1cl.s. Status Suny~ md CorlwnutiorlAc,timPlm. Compiled by D.H.M. Cumming. R.F. du Toit, S.N. Stuart
and the IUCN/ SSC African Elephant and Rhino Specialist Group,
1990. 73 pp. (Out of print.)
Fo.ue.s. Wol~~es, Jtrcktrls, md Dogs. AI/ Actiorl Plall .fi)r-the
Comenutiotr
of’Cmid,s. Compiled by J.R. Ginsberg, D.W.
Macdonald, and the IUCN/SSC
Canid and Wolf Specialist
Groups, 1990, 1 16 pp.
The Asim Elephmt.
A/l 14c*tio/l Plm ,fiw its Comerwtim.
Compiled
by C. Santiapillai, P. Jackson. and the IUCN/SSC Asian ElephantSpecialist Group. 1990, 79 pp.
Allte1ope.s. Glolxrl Sunv~~ md Regiorurl Acatioll PIc11l.s. Part 3.
West md Cmtral,4fricw.
Compiled by R. East and the IUCN/SSC
Antelope Specialist Group, 1990, 17 I pp.
Otters. AI1 Actim Pltrrz ,fi,r their Co~~sen~cltim. Compiled by
P. Foster-Turley.
S. Macdonald, C. Mason and the IUCN/SSC
Specialist Group, 1990, 126 pp.
Rabbits, Hures L/rid Pikm. Status Sun*qt md Comen’atiorl
ActimPlm. Compiled by J.A. Chapman, J.E.C. Flux, and the
IUCN/SSC
Lagomorph Specialist Group, 1990, 168 pp.

Where to order:chequehnternationalcan be obtained

Otter

Diversity~fi-i(w Imec*tilwru ~1~1 Elc’I~h~lrlt-Shr~~~~~s. All Ac*ticm Plcl11.fkr theirC’ollsc~rl~trtiol1. Compiled by M.E. Nicoll, G.B. Rathbun and the
IUCN/SSC Insectivore, Tree-Shrew and Elephant-Shrew
Specialist
Group, 1990, 53 pp.
S\bvlllocvtuil Buttcj@lies. Al1 Ac*tim Plmz .fhr their- Co~l.sc~~-~‘~rtioll.
Compiled by T.R. New, N.M. Collins and the IUCN/SSC
Lepidoptera Specialist Group. 199 1. 36 pp.
Ct-ocodiles. AI1 Ac*tion Plm .f&- their Comun~c~tion. Compiled by
J. Thorbjarnarson,
H. Messel, F.W. King, J.P. Ross and the
IUCN/SSC
Crocodile Specialist Group, 1992, 136 pp.
Seth

Amer-icm Cmzelids. All Acatiorl Plm ,fi)l- their Comenwtio~l.
Compiled by H. Torres and the IUCN/SSC
South American
Camelid Specialist Group, 1992, 58 pp.

AIT Actim Plum .fiw their
Austrulc~sim Mcrrsupitrls am1 Mmotrtwes.
C’orl.s~~~-~~trtio~z.Compiled by M. Kennedy and the IUCN/SSC
Australasian Marsupial and Monotreme
Specialist Group. 1992.
103 pp.
LwIur.s of’hl~u/~~~qw.stu
I: Au Ac*timl PlaIl ,fi,r their Comsenwtio/l:/YY3-IYYY. Compiled by R.A. Mittermeier.
W.R. Konstant. M.E.
Primate Specialist Group,
Nicoll, 0. La ngrand, and the IUCN/SSC
1992, 58 pp. (Out of print.)
Zebras, Asses md Horse.s. AI] Acstim Plarl .fijr th4 Corlscwcrtio~~ c!f’Wild Equids. Compiled by P. Duncan and the IUCN/SSC
Equid
Specialist Group, 1992, 36 pp.
Old World Fruit Buts. Api Acatim PICIP~.fi,r their C~~lzst~n’~rtioll.
Compiled by S. Mickleburgh,
A.M. Hutson, P.A. Racey andthe IUCN/SSC Chiroptera Specialist Group, 1992, 252 pp.
(Out of print.)
Smls, Fur Swls, Sect Lims, md Wcrlrm. Status Sung* md
Corlsc~rlltrtiolz Actim Plm. Peter Reijnders. Sophie Brasseur, Jaap
van der Toorn, Peter van der Wolf, Ian Boyd, John Harwood.
David Lavigne, Lloyd Lowry, and the ILJCN/SSC Seal Specialist
Group, 1993, 88 pp,
Piss, Peccaries, ad HipIms. Status SnnTej* urzd Corzsenwtim
Actim
Pluu. Edited by William L.R. Oliver and the IUCN/SSC
Pigs andPeccaries Specialist Group and the IUCN/SSC
Hippo Specialist
Group, 1993, 202 pp.
The Red PumIa, Olillgo,s, Cmrtis, R~uu~om, md their Rt)luti\*es. Stcrtlrs
Sunq~ clrzd Corwnwtioll
Ac*tim Plm .fijr Proqwids
md Ailw-ids.
(In English and Spanish) Compiled by Angela R Glatston and the
IUCN/SSC
Mustelid, Viverrid, and Procyonid Specialist Group,
1994, 103 pp.
Ikjlphirzs, PorIwi.se.s, crrld Whtrles. I YYd- I YY3 Actim Plm jiw theCor~sc9-v~rtior~ of’Cetatwms.
Compiled by Randall R. Reeves andStephen Leatherwood
together with the IUCN/SSC Cetacean
Specialist Group, 1994, 9 1 pp.
Megq~odes.
AIT Acbtiou Plcm .fijr-their Corlscnwtim
I YY.5- 1YYY.
Compiled by Ren6 W.R.J.Dekker,
Philip J.K.McGowan
and the
WPA/Birdlife/SSC
Megapode Specialist Group, 1995, 4 1 pp.

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