Proceedings of the Seventh Mountain Lion Workshop

PROCEEDINGS OF THE 

SEVENTH MOUNTAIN LION WORKSHOP 

15-17 MAY 2003 • THE VIRGINIAN LODGE • JACKSON, WYOMING 

Editors: 

Scott A. Becker 

Daniel D. Bjornlie 

Fred G. Lindzey 

David S. Moody 

Organizing Committee 

Scott Becker Ron Grogan 

Dan Bjornlie Fred Lindzey 

Tom Easterly Dave Moody 

Sponsored By: 

The Wyoming Chapter of the Wildlife Society 

Wyoming Game and Fish Department 

Wyoming Cooperative Fish and Wildlife Research Unit 

© 2003 


260 Buena Vista 

Lander, Wyoming 82520

Suggested Citation Formats 

Entire Volume: 

Becker, S.A., D.D. Bjornlie, F.G. Lindzey, and D.S. Moody. eds. 2003. Proceedings of the 

Seventh Mountain Lion Workshop. Lander, Wyoming. 

For individual papers: 

Author’s name(s). 2003. Title of Paper. Pages 00-00 in S.A. Becker, D.D. Bjornlie, F.G. 

Lindzey, and D.S. Moody, eds. Proceedings of the Seventh Mountain Lion Workshop. 

Lander, Wyoming. 

Purchasing Additional Copies of the Proceedings 

Please send a check made out to “Wyoming Chapter, TWS” for the amount of fifteen (15) US 

dollars to Tim Thomas, Wyoming Game and Fish Department, PO Box 6249, Sheridan, WY 

82801, USA; phone: (307) 672-7418; email: Tim.Thomas@wgf.state.wy.us. Information on 

different purchasing options may also be made through Tim.

TABLE OF CONTENTS 

Preface....................................................................................................................................................................vii 

In Memory 

Ian Ross ............................................................................................................................................................. viii 

Mountain Lion Status Reports 

Session Chair: Dave Moody, Wyoming Game and Fish Department 

STATUS OF MOUNTAIN LION POPULATIONS IN ARIZONA 

Brian F. Wakeling ....................................................................................................................................................1 

CALIFORNIA MOUNTAIN LION STATUS REPORT 

Doug Updike............................................................................................................................................................6 

COLORADO MOUNTAIN LION STATUS REPORT 

Jerry Apker.............................................................................................................................................................14 

FLORIDA FISH AND WILDLIFE CONSERVATION COMMISSION STATUS REPORT 

Mark Lotz and E. Darrell Land ..............................................................................................................................18 

IDAHO MOUNTAIN LION STATUS REPORT 

Steve Nadeau..........................................................................................................................................................25 

MONTANA MOUNTAIN LION STATUS REPORT 

Rich DeSimone and Rose Jaffe..............................................................................................................................29 

NEVADA MOUNTAIN LION STATUS REPORT 

Russell Woolstenhulme..........................................................................................................................................31 

NEW MEXICO MOUNTAIN LION STATUS REPORT 

Rick Winslow.........................................................................................................................................................39 

STATE OF SOUTH DAKOTA MOUNTAIN LION STATUS REPORT 

Mike Kintigh ..........................................................................................................................................................43 

MOUNTAIN LION STATUS REPORT FOR TEXAS 

John Young ............................................................................................................................................................49 

UTAH MOUNTAIN LION STATUS REPORT 

Craig R. McLaughlin .............................................................................................................................................51 

WASHINGTON COUGAR STATUS REPORT 

Richard A. Beausoleil, Donald A. Martorello, and Rocky D. Spencer..................................................................60 

WYOMING MOUNTAIN LION STATUS REPORT 

Scott A. Becker, Daniel D. Bjornlie, and David S. Moody....................................................................................64 

CRYPTIC COUGARS – PERSPECTIVES ON THE PUMA IN THE EASTERN, MIDWESTERN, AND GREAT PLAINS 

REGIONS OF NORTH AMERICA 

Jay W. Tischendorf ................................................................................................................................................71 

MOUNTAIN LION STATUS REPORT: BRITISH COLUMBIA – Abstract 

Matt Austin ............................................................................................................................................................87 

IMPROVING OUR UNDERSTANDING OF MOUNTAIN LION MANAGEMENT TRENDS: THE VALUE OF CONSISTENT 

MULTI-STATE RECORD KEEPING - Abstract 

Christopher M. Papouchis and Lynn Michelle Cullens .........................................................................................88 

Mountain Lion Interactions with Humans and Livestock 

Session Chair: Kenneth Logan, Colorado Division of Wildlife 

iii

LESSENING THE IMPACT OF A PUMA ATTACK ON A HUMAN 

E. Lee Fitzhugh, Sabine Schmid-Holmes, Marc W. Kenyon, and Kathy Etling ...................................................89 

A CONCEPTUAL MODEL AND APPRAISAL OF EXISTING RESEARCH RELATED TO INTERACTIONS BETWEEN 

HUMANS AND PUMAS – Abstract 

David J. Mattson, Jan V. Hart, Paul Beier, and Jesse Millen-Johnson ................................................................104 

RELATIONSHIPS BETWEEN LAND TENURE SYSTEM, MOUNTAIN LION PROTECTION STATUS, AND LIVESTOCK 

DEPREDATION RATE – Abstract 

Marcelo Mazzolli .................................................................................................................................................105 

MOUNTAIN LION MOVEMENTS AND PERSISTENCE IN A FRAGMENTED, URBAN LANDSCAPE IN SOUTHERN 

CALIFORNIA – Abstract 

Seth P.D. Riley, Raymond M. Sauvajot, and Eric C. York..................................................................................106 

PUMA RESPONSES TO CLOSE ENCOUNTERS WITH RESEARCHERS – Abstract 

Linda L. Sweanor, Kenneth A. Logan, and Maurice G. Hornocker ....................................................................107 

Mountain Lion Genetics and Disease 

Session Chair: Deedra Hawk, Wyoming Game and Fish Department 

PRELIMINARY RESULTS OF FLORIDA PANTHER GENETIC ANALYSES – Abstract 

Warren E. Johnson, Darrell Land, Jan Mortenson, Melody Roelke-Parker, and Stephen J. O’Brien..................108 

GENETIC STRUCTURE OF COUGAR POPULATIONS ACROSS THE WYOMING BASIN: METAPOPULATION OR 

MEGAPOPULATION – Abstract 

Chuck R. Anderson, Jr., Fred G. Lindzey, and Dave B. McDonald ....................................................................109 

ECOLOGICAL SIGNIFICANCE AND EVOLUTION OF A COMMON COUGAR RETROVIRUS – Abstract 

Roman Biek and Mary Poss.................................................................................................................................110 

Mountain Lion Ecology 

Session Chair: Fred Lindzey, Wyoming Cooperative Fish and Wildlife Research Unit 

CHARACTERISTICS OF MOUNTAIN LION BED, CACHE AND KILL SITES IN NORTHEASTERN OREGON 

James J. Akenson, M. Cathy Nowak, Mark G. Henjum, and Gary W. Witmer...................................................111 

IMPACT OF EDGE HABITAT ON HOME RANGE SIZE IN PUMAS – Abstract 

John W. Laundré and Lucina Hernández.............................................................................................................119 

EFFECT OF ROADS ON HABITAT USE BY COUGARS – Abstract 

Dorothy M. Fecske, Jonathan A. Jenks, Frederick G. Lindzey, and Steven L. Griffin........................................120 

ECOLOGY OF SYMPATRIC PUMAS AND JAGUARS IN NORTHWESTERN MEXICO – Abstract 

Carlos A. Lopez Gonzalez and Samia E. Carrillo Percastegui.............................................................................121 

COUGAR ECOLOGY AND COUGAR-WOLF INTERACTIONS IN YELLOWSTONE NATIONAL PARK: A GUILD 

APPROACH TO LARGE CARNIVORE CONSERVATION – Abstract 

Toni K. Ruth, Polly C. Buotte, Howard B. Quigley, and Maurice G. Hornocker................................................122 

EVALUATION OF HABITAT FACTORS THAT AFFECT THE ABUNDANCE OF PUMAS IN THE CHIHUAHUAN 

DESERT – Abstract 

Joel Loredo Salazar, Lucina Hernández, and John W. Laundré ..........................................................................123 

Mountain Lion/Prey Dynamics 

Session Chair: Steve Cain, Grand Teton National Park 

ARE PUMAS OPPORTUNISTIC SCAVENGERS? – Abstract 

Jim W. Bauer, Kenneth A. Logan, Linda L. Sweanor, and Walter M. Boyce .....................................................124 

COUGAR-INDUCED INDIRECT EFFECTS: DOES THE RISK OF PREDATION INFLUENCE UNGULATE FORAGING 

BEHAVIOR ON THE NATIONAL BISON RANGE? – Abstract 

David M. Choate, Gary E. Belovsky, and Michael L. Wolfe ..............................................................................125 

iv

COUGAR PREDATION ON PREY IN YELLOWSTONE NATIONAL PARK: A PRELIMINARY COMPARISON PRE- AND 

POST-WOLF REESTABLISHMENT – Abstract 

Toni K. Ruth, Polly C. Buotte, Kerry M. Murphy, and Maurice G. Hornocker ..................................................126 

FOUR DECADES OF COUGAR-UNGULATE DYNAMICS IN THE CENTRAL IDAHO WILDERNESS – Abstract 

Holly A. Akenson, James J. Akenson, Howard B. Quigley, and Maurice G. Hornocker....................................127 

COUGAR TOTAL PREDATION RESPONSE TO DIFFERING PREY DENSITIES: A PROPOSED EXPERIMENT TO TEST 

THE APPARENT COMPETITION HYPOTHESIS – Abstract 

Hugh Robinson, Robert Wielgus, Hilary Cruickshank, and Catherine Lambert .................................................128 

Mountain Lion Population Monitoring and Management 

Session Chair: Kerry Murphy, Yellowstone National Park 

CHARACTERISTICS OF COUGAR HARVEST WITH AND WITHOUT THE USE OF DOGS 

Donald A. Martorello and Richard A. Beausoleil................................................................................................129 

RESPONSE BY THREE LARGE CARNIVORES TO RECREATIONAL BIG GAME HUNTING ALONG THE YELLOWSTONE 

NATIONAL PARK AND ABSAROKA-BEARTOOTH WILDERNESS BOUNDARY – Presentation Only 

Howard B. Quigley, Toni K. Ruth, Douglas W. Smith, Mark A. Haroldson, Polly C. Buotte, Charles C. 

Schwartz, Steve Cherry, Kerry M. Murphy, Dan Tyers, and Kevin Frey 

DEFINING AND DELINEATING DE FACTO REFUGIA: A PRELIMINARY ANALYSIS OF THE SPATIAL DISTRIBUTION 

OF COUGAR HARVEST IN UTAH AND IMPLICATIONS FOR CONSERVATION – Abstract 

David C. Stoner and Michael L. Wolfe................................................................................................................136 

MONITORING CHANGES IN COUGAR SEX/AGE STRUCTURE WITH CHANGES IN ABUNDANCE AS AN INDEX TO 

POPULATION TREND – Abstract 

Chuck R. Anderson, Jr. and Fred G. Lindzey ......................................................................................................137 

MANAGEMENT OF COUGARS (Puma concolor) IN THE WESTERN UNITED STATES – Abstract 

Deanna Dawn, Michael Kutilek, Rich Hopkins, Sulehka Anand, and Steve Torres ...........................................138 

DYNAMICS AND VIABILITY OF A COUGAR POPULATION IN THE PACIFIC NORTHWEST – Abstract 

Catherine Lambert, Robert B. Wielgus, Hugh S. Robinson, Donald D. Katnik, Hilary Cruickshank, and 

Ross Clarke ..........................................................................................................................................................139 

PROJECT CAT (COUGARS AND TEACHING): INTEGRATING SCIENCE, SCHOOLS AND COMMUNITY IN 

DEVELOPMENT PLANNING – Abstract 

Gary M. Koehler and Evelyn Nelson...................................................................................................................140 

MONITORING MOUNTAIN LIONS IN THE TUCSON MOUNTAIN DISTRICT OF SAGUARO NATIONAL PARK, 

ARIZONA, USING NONINVASIVE TECHNIQUES – Abstract 

Lisa Haynes, Don Swann, and Melanie Culver ...................................................................................................141 

ESTIMATING COUGAR ABUNDANCE USING PROBABILITY SAMPLING: AN EVALUATION OF TRANSECT VERSUS 

BLOCK DESIGN – Abstract 

Chuck R. Anderson, Jr., Fred G. Lindzey, and Nate Nibbelink...........................................................................142 

EVALUATING MOUNTAIN LION MONITORING TECHNIQUES IN THE GARNET MOUNTAINS OF WEST CENTRAL 

MONTANA – Abstract 

Rich DeSimone ....................................................................................................................................................143 

PRESENCE AND MOVEMENTS OF LACTATING AND MATERNAL FEMALE COUGARS: IMPLICATIONS FOR STATE 

HUNTING REGULATIONS – Abstract 

Toni K. Ruth, Kerry M. Murphy, and Polly C. Buotte ........................................................................................144 

Mountain Lion Conservation 

Session Chair: Christopher Papouchis, Mountain Lion Foundation 

MYSTERY, MYTH AND LEGEND: THE POLITICS OF COUGAR MANAGEMENT IN THE NEW MILLENNIUM – 

Abstract 

Rick A. Hopkins...................................................................................................................................................145 

v

RECONCILING SCIENCE AND POLITICS IN PUMA MANAGEMENT IN THE WEST: NEW MEXICO AS A TEMPLATE 

– Abstract 

Kenneth A. Logan, Linda L. Sweanor, and Maurice G. Hornocker ....................................................................146 

COMMUNITY-BASED CONSERVATION OF MOUNTAIN LIONS – Abstract 

Lynn Michelle Cullens and Christopher Papouchis.............................................................................................147 

PUMA MANAGEMENT IN WESTERN NORTH AMERICA: A 100-YEAR RETROSPECTIVE – Abstract 

Steven Torres, Heather Keough, and Deanna Dawn............................................................................................148 

USING COUGARS TO DESIGN A WILDERNESS NETWORK IN CALIFORNIA’S SOUTH COAST ECOREGION – Abstract 

Paul Beier and Kristeen Penrod ...........................................................................................................................149 

MOUNTAIN LIONS AND BIGHORN SHEEP: FACING THE CHALLENGES – Abstract 

Christopher M. Papouchis and John D. Wehausen ..............................................................................................150 

POSTER PRESENTATIONS 

Session Chair: Scott Becker, Wyoming Game and Fish Department 

FACTORS AFFECTING DISPERSAL IN YOUNG MALE PUMAS 

John W. Laundré and Lucina Hernández.............................................................................................................151 

COUGAR EXPLOITATION LEVELS AND LANDSCAPE CONFIGURATIONS: IMPLICATIONS FOR DEMOGRAPHIC 

STRUCTURE AND METAPOPULATION DYNAMICS – Abstract 

David C. Stoner and Michael L. Wolfe................................................................................................................161 

ASSESSING GPS RADIOTELEMETRY RELIABILITY IN COUGAR HABITAT – Abstract 

Trish Griswold, James Briggs, Gary Koehler, and Students at Cle Elum-Roslyn School District ......................162 

USING GPS COLLARS TO DETERMINE COUGAR KILL RATES, ESTIMATE HOME RANGES, AND EXAMINE 

COUGAR-COUGAR INTERACTIONS –Abstract 

Polly C. Buotte and Toni K. Ruth ........................................................................................................................163 

FUNCTIONAL RESPONSE OF COUGARS AND PREY AVAILABILITY IN NORTHEASTERN WASHINGTON – Abstract 

Hilary S. Cruickshank, Hugh S. Robinson, Catherine Lambert, Robert B. Wielgus ...........................................164 

WHAT DOES TEN YEARS (1993-2002) OF MOUNTAIN LION OBSERVATION DATA REVEAL ABOUT MOUNTAIN 

LION-HUMAN INTERACTIONS WITHIN REDWOOD NATIONAL AND STATE PARKS – Abstract 

Gregory W. Holm ................................................................................................................................................165 

DEPREDATION TRENDS IN CALIFORNIA – Abstract 

Sarah Reed, Christopher M. Papouchis, and Lynn Michelle Cullens ..................................................................166 

THE DISTRIBUTION OF PERCEIVED ENCOUNTERS WITH NON-NATIVE CATS IN SOUTH AND WEST WALES, UK: 

RELATIONSHIP TO MODELED HABITAT SUITABILITY – Abstract 

A.B. Smith, F.E. Street Perrott, and T. Hooper....................................................................................................167 

PUMA ACTIVITY AND MOVEMENTS IN A HUMAN-DOMINATED LANDSCAPE: CUYAMACA RANCHO STATE 

PARK AND ADJACENT LANDS IN SOUTHERN CALIFORNIA – Abstract 

Linda L. Sweanor, Kenneth A. Logan, Jim W. Bauer, and Walter M. Boyce .....................................................168 

MODELING OFFSPRING SEX RATIOS AND GROWTH OF COUGARS – Abstract 

Diana M. Ghikas, Martin Jalkotzy, Ian Ross, Ralph Schmidt, and Shane A. Richards .......................................169 

MOUNTAIN LION SURVEY TECHNIQUES IN NORTHERN IDAHO: A THREE-FOLD APPROACH – Abstract 

Craig G. White, Peter Zager, and Lisette Waits...................................................................................................170 

MOUNTAIN LIONS IN SOUTH DAKOTA: RESULTS OF A 2002 PUBLIC OPINION SURVEY – Abstract 

Larry M. Gigliotti, Dorothy M. Fecske, and Jonathan A. Jenks ..........................................................................171 

CRITICAL COUGAR CROSSING AND BAY AREA REGIONAL PLANNING – Abstract 

Michele Korpos....................................................................................................................................................172 

List of Participants............................................................................................................................................173 

vi

PREFACE 

vii 

PREFACE 

The Wyoming Game and Fish Department took great pride in hosting the Seventh 

Mountain Lion Workshop, which was held in conjunction with the Thirty-Ninth North American 

Moose Conference and Workshop and the Fifth Western States and Provinces Deer and Elk 

Workshop. More than 190 people attended the mountain lion workshop representing 27 states, 3 

Canadian provinces, Mexico, Brazil, and the United Kingdom. Numerous state and federal 

agencies, tribal nations, private organizations, academia, and members of the general public were 

represented which attest to the varied and growing interest in mountain lions throughout North 

and South America. 

This workshop would not have been a success without the aid and cooperation of the 

contributors and participants. Financial support, equipment, and manpower provided by the 

Wyoming Chapter of the Wildlife Society, the Wyoming Game and Fish Department, and the 

Wyoming Cooperative Fish and Wildlife Research Unit at the University of Wyoming made this 

workshop possible. A special thanks goes to the members of the organizing committee for their 

aid with all aspects of pre- and post-workshop activities, to the session chairs for keeping the 

workshop moving in a timely fashion, and to the invited speakers who gave thoughtful insight 

into past, present, and future mountain lion management practices and research techniques. 

Many thanks to the Western Association of Fish and Wildlife Agencies (WAFWA) for 

sanctioning the Seventh Mountain Lion Workshop; from this point forward, all mountain lion 

workshops will be sanctioned by WAFWA. 

Finally, we would like to thank all the presenters in the oral and poster sessions for the 

depth of their research and the quality of their presentations. As a result of the efforts you all put 

forth, a standard has been set for presentations at future mountain lion workshops. Keep up the 

great work! 

Scott Becker and Dave Moody 

Workshop Co-Chairs

IN MEMORY 

P. Ian Ross 

Born December 16, 1958 in Goderich, Ontario. 

Died June 29, 2003, age 44, near Nanyuki, Kenya. 

Ian was a true outdoorsman from the beginning, running a trapline while in high school in 

southern Ontario. After graduating from the University of Guelph (1982), his first experiences 

with grizzly bears came in northwestern Alberta, where he studied the impacts of industrial 

development. It was the beginning of an illustrious 20-year career conducting research on large 

mammals in western Canada. 

He worked on cougars in southwestern Alberta from the early 1980’s until 1994. That project 

became one of the longest running research projects on Puma concolor in North America. The 

cougar project received national recognition on radio and television and Ian used that attention to 

foster a thoughtful and effective wildlife conservation message. He participated in the drafting 

of a management plan for cougars in Alberta as well as a conservation strategy for large 

carnivores in Canada. He was the senior author on 9 papers in peer-reviewed journals in 

addition to many other technical reports and popular articles. 

After the cougar project wrapped up, Ian conducted environmental impact studies in western and 

northern Canada. He recently rewrote the grizzly bear status report for COSEWIC. He also 

worked tirelessly with The Wildlife Society-Alberta Chapter dealing with wildlife conservation 

issues. He served as President of the Chapter in 1997. Ian also continued to capture wildlife, 

including grizzly bears, for research projects, and in doing so assisted many graduate students 

with their research. He conducted his capture work using an exacting professional approach 

while retaining an empathy for the wildlife he was pursuing. He cared for each individual and 

did his utmost to conduct captures in a humane manner. 

Ian was a committed and emotional friend and family man. Having no children of his own he 

was a hero to his young nieces, nephews and children of friends. He always remembered 

everyone’s birthdays. He hiked the foothills of the Rockies west of Calgary, as well as the U.S. 

desert southwest, the Canadian Arctic, Mexico and Africa. He loved to hunt and his dinner table 

was a testiment to his hunting prowess. His conservation ethic permeated all of his life. He did 

not consume needlessly and he encouraged all of us to do the same. 

In January 2003, Ian returned to field research when he joined Dr. Laurence Frank on the 

Liakipia Predator Project, a project designed to find ways to allow for the coexistence of hyenas, 

lions, and leopards and people in the agricultural matrix that exists outside national parks in most 

of southern Africa. Two days before his death he was on top of the world having collared his 

first leopard. On the evening he died Ian was tracking a radio-collared lion from a light aircraft. 

Searchers located its wreckage the next morning. Ian Ross died at the peak of his career, doing 

what he loved. 

By 

Martin Jalkotzy 

Arc Wildlife Services 

3527 - 35 Ave. SW 

Calgary, AB, T3E 1A2, Canada 

viii

ix 

IN MEMORY

STATUS OF MOUNTAIN LION POPULATIONS IN ARIZONA 

BRIAN F. WAKELING, Arizona Game and Fish Department, Game Branch, 2221 West 

Greenway Road, Phoenix, AZ 85023 USA 

Abstract: Arizona's mountain lion (Puma concolor) population numbers about 1,000-2,500 animals, and just over 

350 mountain lions were harvested through sport and depredation take in 5 of the last 6 years. Arizona bag limit is 1 

lion per person per year annually, except in a few units where multiple bag limits have been implemented; no 

multiple bag limit has been reached to date. Management for this big game animal is guided by strategic plan, 

species management guidelines, hunt guidelines, and a predation management policy. Management is currently 

under review by an internal team that is examining several predator species, including mountain lions. The internal 

review should be complete by the end of 2003. Public safety incident reports have increased substantially since 

1998. 

INTRODUCTION 

Mountain lion populations within 

Arizona remain robust and are currently 

estimated at 1,000-2,500 despite a prolonged 

drought throughout the southwestern United 

States. Portions of Arizona have received 

record low precipitation during 2002, and 

the decade of the 1990s was the driest on 

records for several portions of Arizona. 

Mule deer (Odocoileus hemionus) 

populations have declined, and in 2003 the 

Arizona Game and Fish Commission 

authorized the lowest number of permits for 

deer hunting since the limited-draw permit 

system was established in Arizona. 

Figure 1. Arizona mountain lion harvest 

trends excluding tribal lands, 1984-2002. 

1 

Proceedings of the Seventh Mountain Lion Workshop 

Mountain lion harvest has remained high, as 

annual statewide harvests have exceeded our 

strategic plan objectives (Arizona Game and 

Fish Department 2001) in 5 of the last 6 

years (Figure 1). 

Mountain lions are classified as big 

game by Arizona statute. Commission order 

has established the bag limit at 1 mountain 

lion per year, except in a few units. 

Successful hunters are required to report 

their harvest within 10 days and answer a 

series of standard questions. Beginning in 

July 2003, hunters will be asked to 

voluntarily provide a tooth, which may be 

used to estimate age through cementum 

annuli and determine gender using genetic 

techniques. The Department is investigating 

making the tooth submission mandatory. 

The management objectives for this species, 

as well as all big game species, are outlined 

in the agency strategic plan, Wildlife 2006 

(Arizona Game and Fish Department 2001) 

and species management guidelines. The 

strategic plan goals, objectives, and speciesspecific 

strategies for mountain lion 

management, that include: 

Objectives 

1. Maintain annual harvest at 250 to 300 

mountain lions (including depredation

2 STATUS OF MOUNTAIN LION POPULATIONS IN ARIZONA · Wakeling 

take). 

2. Provide recreational opportunity for 

3,000 to 6,000 hunters per year. 

3. Maintain existing occupied habitat and 

maintain the present range of mountain 

lions in Arizona. 

Species-Specific Strategies 

1. Maintain a complete database from all 

harvest sources, through a mandatory 

check-out system, including age, sex, 

kill location, etc. to index population 

trend. 

2. Conduct a hunter questionnaire 

biannually. 

3. Evaluate the management implications 

of population and relative density 

estimates. 

4. Implement hunt structures to increase 

and direct harvest emphasis toward 

areas with high lion populations, and 

where depredation complaints are 

substantiated, and evaluate the 

effectiveness of these efforts. 

5. Determine population numbers and 

characteristics on a hunt-area basis. 

6. Increase public awareness of mountain 

lions and their habits, to reduce 

conflicts with humans. 

7. Implement the Department’s Predation 

Management Policy. 

In addition, management direction is 

provided by species management guidelines 

and hunt guidelines. In October 2000, the 

Arizona Game and Fish Commission 

approved the predation management policy 

that provides the agency guidance as to 

when and how to engage in predation 

management. 

Mountain lion management has changed 

as a direct result of biological investigations 

into predation effects. Mountain lion 

predation is being documented as a factor 

PROCEEDINGS OF THE SEVENTH MOUNTAIN LION WORKSHOP 

that may be regulating prey populations 

(Ballard et al. 2001) in some areas of 

Arizona, to include bighorn sheep (Ovis 

canadensis) (Kamler et al. 2002) and 

pronghorn (Antilocapra americana) 

(Ockenfels 1994a, b). These prey 

populations are at low levels, and reducing 

predator populations is likely to allow those 

prey populations to increase in number 

(Ballard et al. 2001). The standard bag limit 

for mountain lions has been altered in 

specific areas to allow for the harvest of 1 

mountain lion per day until a predetermined 

number of mountain lions are removed that 

equal about 50-75% of the estimated 

mountain lion population within that unit, at 

which time the bag limit reverts back to the 

standard bag limit of 1 mountain lion per 

calendar year. The only exception to this is 

in the southwestern portion of the state 

where if even a single mountain lion is 

taken, the hunt area will be closed. 

Multiple bag limits were implemented in 

Units 13A and 13B in 1999, 16A South and 

18B South in 2001, 22 South in 1999, and 

Units 21 West, 28 South, and 37B North 

will be implemented this year. Research 

studies in Unit 22 South on bighorn sheep, 

that included investigations into nutrition, 

disease, and predation, indicate that the 

multiple bag limit on mountain lions in that 

area, with increased effort by sportsmen to 

harvest mountain lions, seems to be 

positively influencing desert bighorn sheep 

recruitment and adult female survival. To 

implement a multiple bag limit on mountain 

lions, biologists must identify a prey species 

that has been reduced due to mountain lion 

predation (e.g., a declining population below 

management objectives) or a management 

action that is likely to be impacted by 

mountain lion predation (e.g., a planned 

translocation) to initiate and identify what 

management objectives must be met (e.g., 3 

years of 50:100 lamb:ewe ratios) before the 

multiple bag limit is removed. Because this

is a relatively recent management approach 

in Arizona, refinements to implementation 

and new opportunities will undoubtedly 

develop. For instance, portions of Arizona 

have robust mountain lion populations that 

sustain large amounts of depredation 

removal (Cunningham et al. 1995) and may 

be able to provide recreational harvest at a 

higher level. These areas might provide 

opportunities to manage recreational harvest 

with multiple bag limits in the future, and 

attempt to transfer depredation take into 

recreational harvest. 

The Department has recently established 

an internal team to review management 

approaches for several predator species, to 

include mountain lions. This team will be 

reviewing social and biological issues and 

best management practices, and 

recommending possible changes to 

Arizona's management. This team will 

serve as an umbrella team for several 

subteams that will work on the biological 

basis for management, gather information on 

social acceptance, and conduct public 

outreach and education. 

DISTRIBUTION AND ABUNDANCE 

Mountain lions are distributed 

throughout most of Arizona, in varying 

densities (Figure 2). This distribution was 

reevaluated in 2002 by Department 

biologists and wildlife managers, and 

although subtle changes have been noted in 

the densities of lions, little change to the 

distribution was identified. This map is still 

undergoing refinement and should be 

considered a draft. Additional information 

used by the Arizona Game and Fish 

Department in managing mountain lion 

population trends includes harvest, 

depredation reports, and age and gender 

from mandatory hunter reports. 

Mountain lion population estimates are 

based on density estimates developed from 

research studies, literature, and professional 

experience within Arizona habitats. These 

STATUS OF MOUNTAIN LION POPULATIONS IN ARIZONA · Wakeling 3 


Figure 2. Mountain lion distribution and 

density estimates (draft) in Arizona 

excluding tribal lands, 2002. 

density estimates are reevaluated at 

infrequent intervals. Prior to 2002, the last 

reevaluation was conducted in 1993, 

although a few management units were 

reevaluated in 1998. 

HARVEST INFORMATION 

Licensed hunters may pursue mountain 

lions in Arizona if they purchase a 

nonpermit tag prior to hunting. The annual 

bag limit is 1 lion, except for areas where a 

multiple bag limit is in place as discussed in 

the introduction. Strategic plan objectives 

for statewide harvests are based on historical 

harvest that removed about 10-15% of the 

estimated statewide population. Recently, 

harvest combined with depredation removal 

has exceeded the strategic plan objective 

(Table 1). Phelps (2003) reported data on 

harvest prior to 1998. Still, statewide 

harvest is probably

4 STATUS OF MOUNTAIN LION POPULATIONS IN ARIZONA · Wakeling 

Table 1. Arizona mountain lion harvest summary excluding tribal lands, 1998-2002. 

Total Harvest 

Sport 

Harvest 


Gender of Sport 

Harvest 

Year 

Tags 

Issued Sport Depredation Other b 

Using Dogs Male Female 

1998 6590 

1999 6885 

2000 7478 

2001 8109 

2002 7900 a 

289 52 1 192 150 136 

247 49 2 161 126 120 

276 53 0 193 133 141 

326 58 0 214 176 144 

263 50 5 175 154 115 

a 

2002 tags sold is preliminary. 

b 

Includes known kills other than sport or depredation (e.g., highway mortality, capture mortality, and illegal take). 

limits are established to take 50-75% of the 

mountain lions that occupy an area when the 

aforementioned criteria are met. To date, 

none of the multiple bag limits have been 

achieved. 

Arizona mountain lion seasons are 

currently open yearlong. About 7,900 

nonpermit tags were sold to hunters in 2002 

(Table 1). During 1998-2002, about 67% 

were taken with the aid of hounds, whereas 

24% were taken incidental to other 

activities. Currently, Arizona does not have 

a pursuit-only season. 

DEPREDATIONS AND HUMAN 

INTERACTIONS-CONFLICTS 

Complaints that come to the Arizona 

Game and Fish Department can take 1 of 2 

routes: nuisance wildlife or depredation. 

Nuisance complaints are dealt with through 

advice and education. Should a mountain 

lion pose a threat to public safety, the 

Department will dispatch a wildlife manager 

to deal with the immediate situation, 

although we frequently contract with USDA 

APHIS Wildlife Services to conduct 

removal efforts. Between 1998 and 2002, 

312 public safety incidents have been 

reported involving mountain lions. The 

trend of these reports over time has been 

steeply increasing (29 in 1998, 105 in 2002; 

Table 2). This increase in reports may be 

Table 2. Public incident reports that included 

mountain lions in Arizona excluding tribal 

lands, 1998-2002. 

Year 

1998 

1999 

2000 

2001 

2002 

Number of Incidents 

Reported 

29 

43 

46 

89 

105 

attributed to mountain lions pursuing prey 

near residential areas (which may be 

exacerbated by drought conditions), 

increasing residential development in 

mountain lion habitat, and the public's 

greater familiarity with the reporting 

process. During that time, few mountain 

lions (

objectives for mountain lions and ranges 

between 49 and 58 mountain lions annually 

(Table 1). The actual number of depredation 

incidents by year is difficult to accurately 

ascertain. 

ONGOING RESEARCH 

Arizona is fortunate to have a Research 

Branch within our Wildlife Management 

Division that may focus on issues 

surrounding wildlife management. In the 

past, this has included many studies directly 

relating to mountain lions and that we 

currently base much of our mountain lion 

management. Currently, the Department 

does not have any ongoing research directly 

aimed at mountain lion management, 

although a study in Unit 22 that includes the 

impacts of mountain lions on desert bighorn 

sheep is being completed. Studies by other 

organizations involving urban mountain 

lions are in the initial phases near Flagstaff 

and Tucson. 

LITERATURE CITED 

ARIZONA GAME AND FISH DEPARTMENT. 

2001. Wildlife 2006. Arizona Game 

and Fish Department, Phoenix. 

BALLARD, W.B., D.L. LUTZ, T.W. KEEGAN, 

L.H. CARPENTER, AND J.C. DEVOS, JR. 

2001. Deer-predator relationships: a 

review of recent North American studies 

with emphasis on mule and black-tailed 

STATUS OF MOUNTAIN LION POPULATIONS IN ARIZONA · Wakeling 5 


deer. Wildlife Society Bulletin 29:99- 

115. 

CUNNINGHAM, S.C., L.A HAYNES, C. 

GUSTAVSON, AND D.D. HAYWOOD. 

1995. Evaluation of the interaction 

between mountain lions and cattle in the 

Aravaipa-Klondyke area of southeast 

Arizona. Arizona Game and Fish 

Department Technical Report 17, 

Phoenix. 

KAMLER, J.F., R.M. LEE, J.C. DEVOS, JR., 

W.B. BALLARD, AND H.A. WHITLAW. 

2002. Survival and cougar predation of 

translocated bighorn sheep in Arizona. 

Journal of Wildlife Management 

66:1267-1272. 

OCKENFELS, R.A. 1994a. Factors affecting 

adult pronghorn mortality rates in central 

Arizona. Arizona Game and Fish 

Department Wildlife Digest Abstract 16, 

Phoenix. 

OCKENFELS, R.A. 1994b. Mountain lion 

predation on pronghorn in central 

Arizona. Southwestern Naturalist 

39:305-306. 

PHELPS, J. 2003. Status report on mountain 

lions in Arizona. Pages 8-10 in L. A. 

Harveson, P. M. Harveson, and R. W. 

Adams, eds. Proceedings of the Sixth 

Mountain Lion Workshop, Texas Parks 

and Wildlife Department, Austin.

CALIFORNIA MOUNTAIN LION STATUS REPORT 

DOUG UPDIKE, Wildlife Programs Branch, California Department of Fish & Game, 1812 9 th 

Street, Sacramento, CA 95814, USA, email: dupdike@dfg.ca.gov 

INTRODUCTION 

California has a statewide mountain lion 

management plan. In 1990, mountain lions 

were legally classified as a “specially 

protected mammal” by the passage of a 

voter initiative (Proposition 117, June 1990 

ballot). Prior to that initiative, lions were 

classified as “game mammals.” 

The objectives for mountain lion 

management in California is to maintain 

healthy, wild populations of mountain lions 

for the benefit and enjoyment of the people 

in the State, to alleviate public safety 

incidents and reduce damage to private 

property (pets and livestock) by mountain 

lions. Mountain lions are not hunted in 

California, and they may be killed only to 

preserve public safety, alleviate damage to 

private property or to protect listed bighorn 

sheep. 

Number 

350 

300 

250 

200 

150 

100 

50 

0 

Mountain Lion Depredation Permits (1972 - 2002) 

6 



Lions are currently distributed 

throughout all suitable habitats within 

California. Lion numbers appear to be 

stable at an estimated 4,000 to 6,000 adults. 

The number of lions in California is 

based upon extrapolating densities 

determined with the use of radio collars. 

These studies have been conducted in 

various locations of the State. The number 

of lions is determined by multiplying the 

densities and the area represented by the 

ecological province. The studies that 

provide local lion density data have been 

conducted over a period of a couple decades. 

Consequently, the Department recognizes 

the estimate has limited application. 

The Department issues depredation 

permits to property owners who have 

experienced damage from a mountain lion 

(Figure 1). 

1972 

1974 

1976 

1978 

1980 

1982 

1984 

1986 

1988 

1990 

1992 

1994 

1996 

1998 

2000 

2002 

Year 

Permits Issued 

Lions Killed 

Figure 1. The number of mountain lion depredation permits issued and the number of 

lions that have been killed as a result in California, 1972-2002.


Mountain lion hunting is prohibited in 

California. No lions have been taken by 

licensed hunters since 1972. It is also illegal 

for lions that have been legally taken in 

other states to be imported into California. 


INTERACTIONS/CONFLICTS 

The Department’s Public Safety 

Guidelines are attached. This policy is 

intended to guide the actions and decisions 

of Department personnel who respond to 

mountain lion incidents. 

A summary of the number of human/lion 

incidents is provided in Table 1. 

We provide educational material to the 

public to foster an understanding and 

appreciation of lions. A recent (May-June 

2000) issue of Outdoor California was 

devoted entirely to mountain lions. Most of 

the articles are viewable at: 

http://www.dfg.ca.gov/coned/ocal/features.html 

In addition, we have produced a 

brochure, “Living with California Mountain 

Lions” which is viewable at: 

http://www.dfg.ca.gov/lion/index.html 

Depredation permits may be issued by 

the Department subject to the conditions 

found in Section 402, California Code of 

Regulations, as follows: 

CALIFORNIA MOUNTAIN LION STATUS REPORT · Updike 7 


a. Revocable permits may be issued by the 

department after receiving a report, from 

any owner or tenant or agent for them, of 

property being damaged or destroyed by 

mountain lion. The department shall 

conduct and complete an investigation 

within 48 hours of receiving such a 

report. Any mountain lion that is 

encountered in the act of inflicting injury 

to, molesting or killing livestock or 

domestic animals may be taken 

immediately if the taking is reported 

within 72 hours to the department and 

the carcass is made available to the 

department. Whenever immediate 

action will assist in the pursuit of the 

particular mountain lion believed to be 

responsible for damage to livestock or 

domestic animals, the department may 

orally authorize the pursuit and take of a 

mountain lion. The department shall 

investigate such incidents and, upon a 

finding that the requirements of this 

regulation have been met, issue a free 

permit for depredation purposes, and 

carcass tag to the person taking such 

mountain lion. 

b. Permittee may take mountain lion in the 

manner specified in the permit, except 

that no mountain lion shall be taken by 

means of poison, leg-hold or metaljawed 

traps and snares. 

Table 1. Summary of the number of human/lion incidents in California, 1995-2002. 

1995 1996 1997 1998 1999 2000 2001 2002 

# of incidents 381 587 539 353 697 372 456 379 

# of safety 

incidents 

18 14 15 11 16 8 14 13 

take 9 7 11 12 10 7 11 13 

male 3 3 1 6 6 4 8 6 

female 3 1 6 6 3 3 3 5 

unknown 3 3 4 0 1 0 0 2 

# of sightings 191 346 340 214 382 174 240 224

8 CALIFORNIA MOUNTAIN LION STATUS REPORT · Updike 

c. Permittee may take mountain lion in the 

manner specified in the permit, except 

that no mountain lion shall be taken by 

means of poison, leg-hold or metaljawed 

traps and snares. 

d. Both males and females may be taken 

during the period of the permit 

irrespective of hours or seasons. 

e. The privilege granted in the permit may 

not be transferred, and only entitles the 

permittee or the employee or agent of 

the permittee to take mountain lion. 

Such person must be 21 years of age or 

over and eligible to purchase a 

California hunting license. 

f. Any person issued a permit pursuant to 

this section shall report by telephone 

within 24 hours the capturing, injuring 

or killing of any mountain lion to an 

office of the department or, if 

telephoning is not practical, in writing 

within five days after capturing, injuring 

or killing of the mountain lion. Any 

mountain lion killed under the permit 

must be tagged with the special tag 

furnished with the permit; both tags must 

be completely filled out and the 

duplicate mailed to the Department of 

Fish and Game, Sacramento, within 5 

days after taking any mountain lion. 

g. The entire carcass shall be transported 

within 5 days to a location agreed upon 

between the issuing officer and the 

permittee, but in no case will a permittee 

be required to deliver a carcass beyond 


the limits of his property unless he is 

willing to do so. The carcass of 

mountain lions taken pursuant to this 

regulation shall become the property of 

the state. 

h. Animals shall be taken in a humane 

manner so as to prevent any undue 

suffering to the animals 

i. The permittee shall take every 

reasonable precaution to prevent the 

carcass from spoiling until disposed of in 

the manner agreed upon under 

subsection (f) of these regulations 

j. The permit does not invalidate any city, 

county, or state firearm regulation. 

k. Permits shall be issued for a period of 10 

days. Permits may be renewed only 

after a finding by the department that 

further damage has occurred or will 

occur unless such permits are renewed. 

The permittee may not begin pursuit of a 

lion more than one mile nor continue 

pursuit beyond a 10-mile radius from the 

location of the reported damage. 

CURRENT RESEARCH 

a. Population genetics of lions 

b. Lion/deer/bighorn sheep predator prey 

relationships in Inyo/Mono counties and 

San Diego County 

c. Lion movements and corridors in Los 

Angeles/Ventura counties 

d. Impacts of habitat conversions and 

transportation corridors or lion 

movements and habitat use.

PUBLIC SAFETY WILDLIFE GUIDELINES – 2072 

Consistent with Section 1801 of the Fish 

and Game Code, these Public Safety 

Wildlife Guidelines provide procedures to 

address public safety wildlife problems. 

Mountain lions, black bears, deer, coyotes, 

and large exotic carnivores that have 

threatened or Attacked humans are wildlife 

classified as public safety problems. Public 

safety wildlife incidents are classified into 

three types: 

A. Type Green (sighting) 

A report (confirmed or unconfirmed) of 

an observation that is perceived to be a 

public safety wildlife problem. The mere 

presence of the wildlife species does not 

in itself constitute a threat. 

B. Type Yellow (threat) 

A report where the presence of the 

public safety wildlife is confirmed by a 

field investigation, and the responding 

person (law enforcement officer or 

Department employee) perceives the 

animal to be an imminent threat to 

public health or safety. Imminent threat 

means there is a likelihood of human 

injury based on the totality of the 

circumstances. 

C. Type Red (attack) 

An attack by a public safety wildlife 

species on a human resulting in physical 

contact, injury, or death. 

These guidelines are not intended to 

address orphaned, injured, or sick wildlife 

that have not threatened public safety. To 

achieve the intent of these guidelines, the 

following procedures shall be used. 

I. Wildlife Incident Report Form 

Fill out a Wildlife Incident Report 

Form (WMD-2) for all reports of public 

safety wildlife incidents. The nature of 

the report will determine the response or 



investigative action to the public safety 

problem. For those reports that require a 

follow-up field investigation, the 

Wildlife Incident Report Form will be 

completed by the field investigator. All 

completed Wildlife Incident Report 

Forms shall be forwarded through the 

regional offices to the Chief, WPB. 

II. Response to Public Safety Wildlife 

Problems 

The steps in responding to a public 

safety wildlife incident are diagramed 

below (Figure 3). 

Any reported imminent threats or 

attacks on humans by wildlife will 

require a follow-up field investigation. 

If a public safety wildlife species is 

outside its natural habitat and in an area 

where it could become a public safety 

problem, and if approved by the Deputy 

Director for the WIFD, it may be 

captured using restraint techniques 

approved by the Wildlife Investigations 

Laboratory (WIL). The disposition of the 

captured wildlife may be coordinated 

with WIL. 

A. Type Green (sighting) 

If the investigator determines that no 

imminent threat to public safety exists, 

Figure 3. Steps in responding to a public 

safety wildlife incident.


the incident is considered a Type Green. 

The appropriate action may include 

providing wildlife behavior information 

and mailing public educational materials 

to the reporting party. 

B. Type Yellow (threat) 

Once the field investigator finds 

evidence of the public safety wildlife 

and perceives the animal to be an 

imminent threat to public health or 

safety, the incident is considered a Type 

Yellow. In the event of threat to public 

safety, any Department employee 

responding to a reported public safety 

incident may take whatever action is 

deemed necessary within the scope of 

the employee's authority to protect 

public safety. When evidence shows that 

a wild animal is an imminent threat to 

public safety, that wild animal shall be 

humanely euthanized (shot, killed, 

dispatched, destroyed, etc.). For Type 

Yellow incidents the following steps 

should be taken: 

1. Initiate the Incident Command 

System. The Incident Commander 

(IC) consults with the regional 

manager or designee to decide on the 

notification process on a case-bycase 

basis. Full notification includes: 

the field investigator's supervisor, the 

appropriate regional manager, the 

Deputy Director, WIFD, Chief, 

Conservation Education and 

Enforcement Branch (CEEB), Chief, 

WPB, WIL, Wildlife Forensics Lab 

(WFL), the designated regional 

information officer, and the local law 

enforcement agency. 

2. If full notification is appropriate, 

notify Sacramento Dispatch at (916) 

445-0045. Dispatch shall notify the 

above-mentioned personnel. 


3. Secure the scene as appropriate. 

Take all practical steps to preserve 

potential evidence. The IC holds 

initial responsibility and authority 

over the scene, locating the animal, 

its resultant carcass, and any other 

physical evidence from the attack. 

The IC will ensure proper transfer 

and disposition of all physical 

evidence. 

4. In most situations, it is important to 

locate the offending animal as soon 

as practical. WIL may be of 

assistance. The services of USDA, 

Wildlife Services (WS) can be 

arranged by the regional manager or 

designee contacting the local WS 

District Supervisor. If possible, avoid 

shooting the animal in the head to 

preserve evidence. 

5. If an animal is killed, the IC will 

decide on the notification process 

and notify Sacramento Dispatch if 

appropriate. Use clean protective 

gloves while handling the carcass. 

Place the carcass inside a protective 

durable body bag (avoid dragging 

the carcass, if possible). 

C. Type Red (attack) 

In the event of an attack, the 

responding Department employee may 

take any action necessary that is within 

the scope of the employee's authority to 

protect public safety. When evidence 

shows that a wild animal has made an 

unprovoked attack on a human, that wild 

animal shall be humanely euthanized 

(shot, killed, dispatched, destroyed, etc.). 

For Type Red incidents the following 

steps should be taken: 

1. Ensure proper medical aid for the 

victim. Identify the victim (obtain 

the following, but not limited to: 

name, address, phone number).

2. Notify Sacramento Dispatch at (916) 

445-0045. Dispatch shall notify the 

field investigator's supervisor, the 

appropriate regional manager, the 

Deputy Director, WIFD, Chief, 

CEEB, Chief, WPB, WIL, WFL, the 

designated regional information 

officer, and the local law 

enforcement agency. 

3. Initiate the Incident Command 

System. If a human death has 

occurred, an Enforcement Branch 

supervisor or specialist will respond 

to the Incident Command Post and 

assume the IC responsibilities. The 

IC holds initial responsibility and 

authority over the scene, locating the 

animal, its resultant carcass, and any 

other physical evidence from the 

attack. The IC will ensure proper 

transfer and disposition of all 

physical evidence. 

4. Secure the area as needed. Treat the 

area as a crime scene. In order to 

expedite the capture of the offending 

animal and preserve as much onscene 

evidence as possible, the area 

of the incident must be secured 

immediately by the initial responding 

officer. The area should be excluded 

from public access by use of flagging 

tape or similar tape (e.g., "Do Not 

Enter") utilized at crime scenes by 

local law enforcement agencies. One 

entry and exit port should be 

established. Only essential 

authorized personnel should be 

permitted in the excluded area. A 

second area outside the area of the 

incident should be established as the 

command post. 

5. In cases involving a human death, 

WFL personnel will direct the 

gathering of evidence. Secure items 

such as clothing, tents, sleeping bags, 



objects used for defense during the 

attack, objects chewed on by the 

animal, or any other materials which 

may possess the attacking animal's 

saliva, hair, or blood. 

6. If the victim is alive, advise the 

attending medical personnel about 

the Carnivore Attack-Victim 

Sampling Kit for collecting possible 

animal saliva stains or hair that 

might still be on the victim. If the 

victim is dead, advise the medical 

examiner of this evidence need. This 

sampling kit may be obtained from 

the WFL. 

7. It is essential to locate the offending 

animal as soon as practical. WIL 

may be of assistance. The services of 

WS can be arranged by the regional 

manager or designee contacting the 

local WS District Supervisor. If 

possible, avoid shooting the animal 

in the head to preserve evidence. 

8. If an animal is killed, the IC will 

notify Sacramento Dispatch. Treat 

the carcass as evidence. Use clean 

protective gloves and (if possible) a 

facemask while handling the carcass. 

Be guided by the need to protect the 

animal's external body from: loss of 

bloodstains or other such physical 

evidence originating from the victim; 

contamination by the animal's own 

blood; and contamination by the 

human handler's hair, sweat, saliva, 

skin cells, etc. Tape paper bags over 

the head and paws, then tape plastic 

bags over the paper bags. Plug 

wounds with tight gauze to minimize 

contamination of the animal with its 

own blood. Place the carcass inside a 

protective durable body bag (avoid 

dragging the carcass, if possible). 

9. WFL will receive from the IC and/or 

directly obtain all pertinent physical


evidence concerning the primary 

questions of authenticity of the 

attack and identity of the offending 

animal. WFL has first access and 

authority over the carcass after the 

IC. WFL will immediately contact 

and coordinate with the county 

health department the acquisition of 

appropriate samples for rabies 

testing. Once WFL has secured the 

necessary forensic samples, they will 

then release authority over the 

carcass to WIL for disease studies. 

10. An independent diagnostic 

laboratory approved by WIL will 

conduct necropsy and disease studies 

on the carcass. The WIL will retain 

primary authority over this aspect of 

the carcass. 

D. Responsibilities of WIL 

WIL investigates wildlife disease 

problems statewide and provides 

information on the occurrence of both 

enzootic and epizootic disease in 

wildlife populations. Specimens 

involved in suspected disease problems 

are submitted to WIL for necropsy and 

disease studies. Most animals killed for 

public safety reasons will be necropsied 

to assess the status of health and whether 

the presence of disease may have caused 

the aggressive and/or unusual behavior. 

Type Yellow public safety animals 

killed may be necropsied by WIL or an 

independent diagnostic laboratory 

approved by WIL. Contact WIL 

immediately after a public safety animal 

is killed to determine where it will be 

necropsied. Arrangements are to be 

made directly with WIL prior to 

submission of the carcass to any 

laboratory. 

Type Red public safety animals 

killed will be necropsied by an 

independent diagnostic laboratory 


approved by WIL. Contact WIL prior to 

submission of the carcass to any 

laboratory to allow the Department 

veterinarian to discuss the disease testing 

requirements with the attending 

pathologist. A disease testing protocol 

has been developed for use with Type 

Red public safety wildlife. 

E. Responsibilities of WFL 

WFL has the statewide responsibility 

to receive, collect, examine and analyze 

physical evidence, issue reports on 

evidence findings, and testify in court as 

to those results. WFL's primary 

functions in public safety incidents is to 

verify or refute the authenticity of the 

purported attack and to corroborate or 

refute the involvement of the suspected 

offending animal. 

Type Yellow public safety animals 

killed may be examined by WFL 

personnel. The examination of the 

carcass will be coordinated with WIL. 

All Type Red public safety animals 

killed must be examined by WFL 

personnel or a qualified person approved 

by WFL supervisor using specific 

procedures established by WFL. 

If a human death occurs, 

coordination of the autopsy between the 

proper officials and WFL is important so 

that WFL personnel can be present 

during the autopsy for appropriate 

sampling and examination. In the event 

of human injury, it is important for WFL 

to gather any relevant physical evidence 

that may corroborate the authenticity of 

a wildlife attack, prior to the treatment of 

injuries, if practical. If not practical, 

directions for sampling may be given 

over the telephone to the emergency 

room doctor by WFL. 

F. Media Contact 

Public safety wildlife incidents

attract significant media attention. Issues 

regarding site access, information 

dissemination, the public's safety, 

carcass viewing and requests to survey 

the scene can be handled by a designated 

employee. Each region shall designate 

an employee with necessary ICS training 

to respond as a regional information 

officer to public safety wildlife 

incidents. 

Type Yellow public safety wildlife 

incidents may require the notification of 

a designated employee previously 

approved by the regional manager or 

designee to assist the IC in responding to 



the media and disseminating 

information. The IC has the authority to 

decide if the designated employee 

should be dispatched to the site. 

All Type Red public safety wildlife 

incidents require that a designated 

employee, previously approved by the 

regional manager or designee, to assist 

the IC in responding to the media and 

disseminating information, is called to 

the scene. 

The Department will develop and 

provide training for designated 

employees to serve as information 

officers for public safety wildlife 

incidents.

COLORADO MOUNTAIN LION STATUS REPORT 

JERRY A. APKER, Colorado Division of Wildlife, 0722 South Road 1 East, Monte Vista, CO 

81144, USA, email: jerry.apker@state.co.us 

MOUNTAIN LION CLASSIFICATION 

Mountain lion (Puma concolor) received 

no legal protection and were classified as a 

predator in Colorado from 1881 until 1965. 

During this time take of puma at any time, 

any place was encouraged by bounties and 

other laws. The first bounty was enacted in 

1881 at $10, in 1925 laws instructed game 

wardens to destroy predatory animals by 

trapping, poisoning, or hunting, and in 1929 

the bounty was increased to $50. For 

comparison the 1929 bounty, if offered in 

2003 dollars, would be $540. The bounty 

was abolished in 1965, but some provisions 

for landowner take of a depredating puma 

remains in Colorado laws to this day. In 

1965, puma were reclassified as big game. 

Each Data Analysis Unit (DAU) within 

the State has a management plan developed 

with objectives for hunter harvest, game 

damage, and human-puma conflicts. 

Objectives are stated as the maximum level 

on a three-year running average. 

Implementation of DAU plans began in 

2001. Recent interest in annual puma kill 

revealed conflicting direction depending 

upon which objectives managers weighed 

most heavily. These conflicts pointed out a 

shortfall within the plans in that they do not 

state a specific strategic goal for the DAU. 

Currently this must be inferred in the text of 

the plan. Some DAUs are managed to 

suppress puma populations while others are 

managed to maintain stable populations – 

recognizing the inherent difficulty in 

determining population changes. Within the 

next year all management plans will be 

required to develop a strategic goal. We 

14 


consider this an essential step for informing 

management decisions within a DAU about 

season structure and annual license 

allocation. 

In 1996 the Colorado Department of 

Agriculture (CDA) was granted “exclusive 

jurisdiction over the control of depredating 

animals that pose a threat to an agricultural 

product or resource”. Thus, CDA has 

exclusive authority to determine the 

disposition of an individual puma if it is 

depredating on livestock, while the Colorado 

Division of Wildlife (CDOW) retains 

authority to manage puma populations and 

all forms of recreational or scientific use. 

DISTRIBUTION, ABUNDANCE AND 

MONITORING 

The state is divided into 21 DAUs for 

the purposes of puma management (Figure 

1). DAUs are assemblages of Game 

Management Units (GMUs) within which 

Figure 1. Data analysis Units and relative 

abundance of puma within each DAU in 

Colorado.

Figure 2. Areas of puma occupancy in 

Colorado. 

puma occupancy has been mapped (Figure 

2). 

Colorado does not regularly estimate 

puma populations because no reliable, cost 

effective sample based population 

estimation technique currently exists. A 

projection of possible population has been 

made based on densities reported in 

literature for intensively studied populations. 

Low and high densities were selected from 

study areas that had habitat types most 

similar to Colorado. Densities were then 

applied by biologists to area of puma habitat 

within DAUs. Areas not considered puma 

habitats, such as extreme high elevations, 

intensively farmed land, cities, highways, or 

reservoirs, were first deleted. Biologists 

were allowed to apply more constrained 

densities based upon their knowledge of 

prey abundance or relative puma abundance. 

Finally, biologists were asked to pinpoint 

the puma density most applicable to DAUs 

within their management responsibility. 

These exercises resulted in a crude projected 

puma population of 3,000 to 7,000, with 

3,500 to 4,500 most probable. Based upon 

the foregoing, each DAU is assigned a 

relative abundance rating of high, moderate, 

or low with intergrades where estimated 

puma density is close to break points. High 

COLORADO MOUNTAIN LION STATUS REPORT · Apker 15 


abundance is assigned at DAU densities of 

over 3 puma/100 km 2 , moderate abundance 

at 2 to 3 puma/100 km 2 , and low abundance 

at anything less than 2 puma/100 km 2 

(Figure 1). 

Hunter harvest and total mortality is 

examined at the DAU level to monitor 

mortality for crude indications of population 

change. Puma mortality is documented 

through mandatory checks of hunter kill and 

mandatory reports for non-hunter mortality 

and is kept in a database. The database for 

hunter kill has been kept since 1980, and for 

non-hunter mortality since 1991. Mortality 

data is examined on three and ten year 

running averages due to relatively high 

annual variation. Data on depredation 

claims since is also maintained in a 

database. 

HARVEST AND HUNTING 

REGULATION 

Since 1972 a quota system has been used 

to manage hunter distribution and kill. From 

1992 the quota has increased from 459 to 

790 in 2002. However, the quota does not 

represent the harvest objective since the 

quota is never achieved. Through 

compilation of DAU management plan 

objectives the harvest objective for the state 

is about 350 puma. Annual license sales 

have also increased since 1992 from about 

900 to just over 1,700 in 2002. While both 

quotas and license sales have increased over 

the past 10 years, percent of quota 

achievement and success relative to license 

sales have declined gradually (Figure 3). 

These trends are expected with increased 

available hunting opportunity toward a 

cryptic species. With more potential hunters 

there is an increased likelihood that there 

will be proportionately more hunters with 

less experience and less commitment or 

impetus to harvest an animal. Some have 

speculated that the trends indicate that overharvest 

has occurred, however the female

16 COLORADO MOUNTAIN LION STATUS REPORT · Apker 

Percent Q. Achievement & Success 

90% 

80% 

70% 

60% 

50% 

40% 

30% 

20% 

10% 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 

% Quota Achievement % Harvest Success by licenses sold 

Quota # Licenses Sold 

1700 

1500 

1300 

1100 

Figure 3. Colorado license sales, quota, 

percent success and percent quota 

achievement for puma. 

component of hunter harvest has not 

increased substantially which would be an 

indicator of over-harvest. 

Criteria used to guide quota setting are 

as follows: 

1. Strategic objective of the DAU or 

group of GMUs within a DAU. If 

management is directed at 

maintaining a stable population, then 

the following also apply. 

2. Population for the DAU is projected 

based upon low and high density 

potential. Off-take should not 

exceed a bracketed range of 15% of 

low-end population estimate and 8% 

of high-end population estimate. 

3. Short (3 year) and long-term trend 

(10 years) in proportion of females in 

mortality should be stable or 

downward and not over 50%. 

4. Damage claim amounts on 3-year 

average should not exceed DAU 

objective levels. 

5. Catch per unit effort indice (effort of 

houndsmen to harvest). 

900 

700 

500 

300 

100 

Quota or License #'s 


Hunter harvest and total mortality 

figures for 2002 have not been completely 

tabulated at the time of this report. The 

average hunter harvest from 1992-1994 is 

308 with 41% female. The average hunter 

harvest from 1999-2001 is 365 with 45% 

female (Figure 4). 

Generally, from 1965 to the mid-late 

1970s seasons were mid fall through early 

spring. In the late 1970s through 1994 

seasons were liberalized, running almost 

continually through the year excluding late 

August – mid November deer or elk hunting 

seasons. Since 1995, seasons were revised 

to provide greater protection for pregnant 

females or females with dependent young, 

running on a calendar year basis from 

January 1 – March 31 and mid November – 

December 31. With a few exceptions the 

bag limit has remained 1 per year of either 

sex and some form of puma license has been 

required since 1965. 

Hunting with hounds is permitted with 

hunting pack size limited to 8 dogs. Almost 

all puma are harvested with the use of 

hounds. There is no pursuit only season. 

With certain technical restrictions on each, 

legal weapons for take include rifle, 

500 

450 

400 

350 

300 

250 

200 

150 

100 

50 

0 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 

Hunter Harvest 

- Male 

Hunter Harvest 

- Female 

Total 

Mortality 

Figure 4. Puma harvest and total mortality 

levels in Colorado.

handgun, shotgun, muzzleloading rifles, 

hand-held bows, and crossbows. It is illegal 

to kill a kitten or a female accompanied by 

kittens. 

DEPREDATION AND PUMA-HUMAN 

CONFLICT 

Colorado is liable for damage caused by 

big game, with certain limitations and 

restrictions. From 1972 until 2001 CDOW 

had to pay for damage by puma and black 

bear to any real or personal property. Black 

bear damage claims often included vehicles, 

buildings, appliances, etc., as well as 

livestock, but puma damage claims have 

been restricted to cattle, sheep, or other 

animals. Beginning in 2001, State liability 

was limited to agricultural products and 

property used in the production of raw 

agricultural products. Liability was also 

changed so that the State is not liable for 

more than $5,000 per animal. 

With the exception of 2000 the number 

of damage claims and the cost of damage 

have declined since 1997 (Figure 5). High 

damage costs in 2000 were mostly due to 6 

claims for the loss of 8 exotic domestic 

animals such as alpaca, llama, and 

250000 

225000 

200000 

175000 

150000 

125000 

100000 

75000 

50000 

25000 

0 

1979 

1981 

1983 

1985 

1987 

1989 

1991 

1993 

1995 

Sheep Cattle Other Stock 

Figure 5. Amount paid on claims for 

depredation by puma in Colorado. 

1997 

1999 

2001 

COLORADO MOUNTAIN LION STATUS REPORT · Apker 17 


commercially owned elk. Procedures for 

handling damage claims are governed by 

statute, regulations, and a game damage 

procedures manual. 

The State has no specific policy 

document providing direction for handling 

puma-human conflicts. However, following 

a human fatality in 1991, DOW staff 

developed procedures that have generally 

been adopted. Encounters involving puma 

are categorized as sightings, encounter 

involving pets, aggressive behavior toward 

humans, or attack on humans. Agency 

responses to these types of encounters vary 

from providing education and information to 

pursue-kill the puma. In the past 5 years, 

fewer than 5-10 encounters beyond sightings 

are documented each year. 

On average over the past 5 years about 

20 puma per year are killed for reasons other 

than hunting. Most of these, 12 per year, are 

control actions on depredating animals. The 

remainders are the result of road kills or 

illegal kills. Less than 1 per year on average 

are killed due to human safety concerns. 

PUMA RESEARCH PROGRAMS 

There are no current research 

investigations being conducted on puma. 

The Division of Wildlife is in the process of 

hiring a research scientist specializing in 

carnivores with emphasis on puma initially.

FLORIDA FISH AND WILDLIFE CONSERVATION COMMISSION STATUS REPORT 

MARK LOTZ, Panther Section Biologist, Florida Fish and Wildlife Conservation Commission, 

566 Commercial Blvd., Naples, FL 34104-4709, USA, email: Mark.Lotz@fwc.state.fl.us 

E. DARRELL LAND, Panther Section Leader, Florida Fish and Wildlife Conservation 

Commission, 566 Commercial Blvd., Naples, FL 34104-4709, USA, email: 

Darrell.Land@fwc.state.fl.us 

INTRODUCTION 

The Florida panther (Puma concolor 

coryi) has been classified as endangered by 

the state of Florida since 1958 and by the 

federal government since 1967. Formerly, 

panthers inhabited the southeastern United 

States, ranging from southern Florida to 

Arkansas and northward to Tennessee and 

South Carolina. Loss and fragmentation of 

habitat coupled with unregulated killing 

over the past two centuries have reduced and 

isolated the panther to the point where only 

one population exists on approximately 

8,810 km 2 of habitat in south Florida (Maehr 

1990). The Florida Fish and Wildlife 

Conservation Commission (FWC) and the 

U. S. Fish and Wildlife Service (USFWS) 

are the two lead authorities involved in all 

aspects of Florida panther recovery and 

protection. Other agencies involved in 

panther recovery include the Florida 

Department of Environmental Protection, 

Florida Division of Forestry, National Park 

Service, South Florida Water Management 

District, as well as numerous nongovernmental 

organizations such as Florida 

Wildlife Federation, National Wildlife 

Federation, The Nature Conservancy, and 

the Florida Audubon Society. A recovery 

plan for the Florida panther was written in 

1981 with revisions in 1987 and 1995 with 

the objective of achieving three viable selfsustaining 

populations within the historic 

range. FWC initiated intensive research 

efforts in 1981 and these studies continue 

18 


today. By the end of 2002, FWC has 

handled 115 panthers for radio-telemetry 

studies and marked 142 neonate kittens at 

dens. FWC and many collaborators have 

published more than 200 papers and reports 

detailing panther life history, habitat use, 

mortality, dispersal, home range dynamics, 

biomedical findings, genetics, population 

modeling, and food habits. 

Florida panthers are threatened by 

demographic instability inherent in small, 

geographically isolated populations and 

erosion of genetic diversity from restricted 

gene flow and inbreeding. Genetic diversity 

is the basis for production of fit individuals 

as well as providing population elasticity in 

order to respond to changing environmental 

and habitat conditions. Historically, natural 

exchange of genetic material occurred 

among the Florida panther population in the 

southeastern United States and contiguous 

populations of P. c. cougar to the north, P. 

c. hippolestes to the northwest and P. c. 

stanleyana to the west (Young and Goldman 

1946). Genetic exchange between 

populations ceased as the coastal plain was 

gradually cleared and settled. Florida 

panthers steadily declined in abundance and 

distribution as a result. Inbreeding increased 

when potential breeders could no longer 

move among fragmented populations and 

the declining population size compounded 

demographic and genetic factors. A 

population viability analysis was conducted 

in 1992, which predicted the extinction of

the Florida panther within 24-63 years (Seal 

1992) and lead to the creation of A Plan for 

Genetic Restoration and Management of the 

Florida Panther (Seal 1994). 

Genetic restoration of the Florida 

panther was implemented in 1995 with the 

release of 8 female Texas cougars (P. c. 

stanleyana) into areas occupied by Florida 

panthers. Five of the 8 cougars produced a 

total of 20 offspring and many of these 

offspring have survived and reproduced. 

The genetic restoration plan identified a goal 

of incorporating a 20% introgression of 

Texas puma genes into the panther 

population and a preliminary assessment 

suggested that we may have achieved or 

slightly exceeded that level (Land and Lacy 

2000). As of January 2003, 5 of the original 

8 released Texas puma have since died and 

the remaining 3 females, thought to be 

reproductively senescent, were removed 

from the wild. We will continue monitoring 

panther genetic restoration by comparing 

reproductive performance, survival, 

phenotypic traits, and genetic characteristics 

among Texas and Florida descendants. Our 

goal is to develop a long-term management 

plan based on our study results to maintain 

genetic diversity, health, and long-term 

survival of the south Florida panther 

population. 


Florida panthers occupy a core range in 

south Florida primarily in Collier, Hendry, 

Lee, and Dade counties. Major public lands 

include Big Cypress National Preserve, 

Everglades National Park, Florida Panther 

National Wildlife Refuge, Fakahatchee 

Strand State Preserve, Picayune Strand State 

Forest, and Okaloacoochee Slough State 

Forest. Large privately held ranches, used 

primarily for cattle and crop production, also 

constitute some of the most important 

habitat for panthers. Verified evidence, 

through road-kills, photos, or tracks, has 

also been found in Glades, Sarasota, and 


FLORIDA STATUS REPORT · Lotz and Land 19 

Palm Beach Counties within the past 2 years 

(Land et al. 2002, Shindle et al. 2001). 

However, these have all been dispersed or 

transient males. No females have been 

documented outside of the core range. One 

radio-collared male panther dispersed a 

straight-line distance of 224 km from his 

natal range (Maehr et al. 2002). 

The first Florida panther was radiocollared 

in 1981 by the Florida Game and 

Fresh Water Fish Commission (renamed to 

Florida Fish and Wildlife Conservation 

Commission in 2000). Throughout the 

1980’s the population was estimated to be 

30-50 adults. The population has been 

increasing since about the mid 1990’s and 

today is estimated to be 80-100 adults. The 

release of Texas cougars for genetic 

restoration purposes in 1995 has contributed 

to this increase. Our population estimate is 

derived by counting currently radio-collared 

panthers and tallying observations of 

uncollared panther sign encountered during 

yearly field activities. 


CONFLICTS 

FWC does not have a specific panther 

depredation or other human conflict protocol 

in place, but we do have a nuisance black 

bear policy that could provide guidance. 

The nuisance bear policy involves 

addressing the source of the problem, 

typically the removal or protection of bear 

attractants, prior to any stepwise progression 

of capture/handling of bears, removals, and 

ultimately, euthanasia. There have been no 

documented panther attacks on humans in 

Florida with only anecdotal accounts of 

attacks prior to 1900 (Tinsley 1970). FWC 

regularly receives complaints about wildlife 

attacks on domestic livestock, many of 

which are claimed to be panther 

depredations. However, upon investigation, 

the vast majority of these incidents involve 

other predators including black bear, bobcat, 

fox, raccoon, opossum, coyote, and

20 FLORIDA STATUS REPORT · Lotz and Land 

domestic dog. We are aware of three valid 

panther depredations that were reported to 

FWC. The first involved a panther that 

seized a small dog by the head and 

subsequently dropped the dog alive after the 

owner appeared at the door. A second 

depredation involved the killing of small 

goats from a rural homeowner’s yard in an 

area occupied by panthers. These 

complainants were given advice on how to 

protect their pets/livestock and to date no 

further depredations have been reported. 

The last case was more complicated because 

it involved panthers that were taking 

advantage of a hunting preserve that was 

newly created by the Seminole Tribe on 

tribal lands. Non-native ungulates were 

stocked in an area known to be occupied by 

panthers and predictably, the panthers 

preyed upon these ungulates. FWC and the 

USFWS could do very little to address these 

depredations because of the Endangered 

Species Act and because the preserve was 

developed on areas used by panthers. 

Although the tribe made a request for 

reimbursement of losses, no compensation 

was provided. Over time, the Seminole 

Tribe has adjusted the type of game animals 

that are stocked in the preserve, primarily 

stocking and selling wild hog hunts, and 

these lower cost animals that are taken by 

panthers are less of a financial loss than the 

various exotic deer species they once 

stocked. Cattle ranchers apparently are 

unconcerned about potential panther 

depredations based on the lack of 

complaints, and FWC food habits work has 

revealed that cattle are rarely taken by 

panthers. The presence of feral hogs on 

cattle ranches provide an abundant, easily 

taken prey base that may obviate the need 

for panthers to tackle cattle. 

RESEARCH AND PUBLICATIONS 

Current Research 


Florida Panther Genetic Restoration and 

Management 

This has been our focal study since 1995 

when 8 female Texas cougars were 

released to offset the problems of 

inbreeding. Genetic diversity and health 

of the Florida panther population needs 

to be restored to ensure survival, even 

with adequate habitat conservation and 

other enhancement measures. Genetic 

restoration is a direct and immediate 

action that will restore genetic variability 

and vitality for a healthier, more resilient 

population. The Plan for Genetic 

Restoration and Management of the 

Florida Panther (Seal 1994) called for a 

20% introgression level of Texas genes 

throughout the population and 

preliminary analysis indicates we are on 

target. All Texas females have died or 

been removed. A minimum of 59 

intercross animals were produced and it 

is assumed that 44 still exist within the 

population. Fifteen are radio-collared. 

This study was extended in order to 

collect and analyze critical samples from 

subsequent generations of Texas puma 

descendants. Our goal is to develop a 

long-term management plan based on 

our study results to maintain genetic 

diversity, health, and long-term survival 

of the south Florida panther population. 

A final report is anticipated next year. 

Feasibility of Using GPS Radio-collars 

on Florida Panthers 

The use of GPS technology in wildlife 

applications has garnered much interest 

in recent years but the current state of 

the technology and its applicability to 

panthers has yet to be determined. 

Among the objectives of this study are to 

compare and evaluate GPS and aerial 

telemetry relocations, calculate the 

percentage of successful GPS 

relocations, and evaluate the use of GPS 

collars on Florida panthers and make

ecommendations for future use. We 

placed 4 GPS collars from Telemetry 

Solutions (1130 Burnett Avenue, Suite J, 

Concord, CA 94520) on panthers during 

our 2001-2002 capture season. Two 

were Posrec collars that stored data on 

board until the collar was retrieved and 

the other two were Simplex units that 

had the ability to transmit data for 

remote downloads as well as store-onboard 

capabilities. Additionally, each 

collar was equipped with a VHF beacon 

in order to detect and recover carcasses, 

pinpoint and visit dens, and enable 

comparisons between GPS locations and 

aerial VHF relocations. Each panther 

equipped with a GPS collar was located 

thrice weekly during our regularly 

scheduled telemetry flights. All GPS 

collars have been recovered and we are 

currently evaluating data and 

performance. Two panthers wearing 

Posrec collars died 7 months after 

deployment, one Simplex model 

failed completely after only 4 months, 

and the remaining Simplex’s main 

battery failed after 6 months, disrupting 

GPS capabilities, but VHF function was 

maintained through the back-up battery. 

A final report is scheduled to be 

completed by the end of 2003. 

Feasibility of Using Remote Cameras to 

Survey Florida Panthers 

Most of what is known about Florida 

panthers, including population 

demographics, has come from radiotelemetry 

studies over the past 20 years. 

However, standardized survey 

techniques that estimate panther 

population parameters with associated 

measures of statistical confidence and 

that document significant changes in 

these parameters over time have not 

been applied. The objective of this study 

is to assess whether infrared-triggered 

camera surveys for panthers provide 



adequate data for inclusion into capturerecapture 

models based on the Lincoln- 

Peterson estimator. Remote camera 

surveys could complement existing labor 

and cost-intensive survey methodology 

to provide a more accurate estimate of 

panther population parameters and 

document significant changes in these 

parameters over time. Passive infrared 

cameras (Cam Trakker, CamTrak 

South Inc., Watkinsville, GA) were 

deployed systematically on two areas 

within the current occupied range of 

Florida panthers. The Florida Panther 

National Wildlife Refuge provided an 

opportunity to assess camera survey 

methodology in a core area with a 

sample population of radio-collared and 

uncollared panthers. Long Pine Key 

within Everglades National Park 

provided an opportunity to assess 

camera survey methodology in a quasigeographically 

closed population of 

radio-collared and uncollared panthers. 

Cameras were systematically placed in 

each study area and trials of 15 and 30 

days were run with 15 and 30 cameras 

per session. Field work was completed 

in 2002 and a final report is expected 

later this year. 

Feasibility of Extracting Florida Panther 

DNA from Scats 

Panther scats could potentially offer the 

safest and most cost effective tool for 

censussing numbers of panthers, 

measuring population genetic health, and 

identifying origins of Puma sign found 

outside of core panther areas. The 

purpose of this study is to evaluate the 

use of panther scats as a source of DNA 

samples for on-going genetic 

monitoring. Existing tissue samples 

were used to calibrate and verify the 

utility of extracting and analyzing DNA 

from scats. Scat collection routes were 

established along existing trails on four


areas and regularly surveyed by ATV. 

Over 400 km of trail were surveyed with 

scats encountered every 45 km on 

average. Additionally, scats were 

collected opportunistically during other 

field activities. Results are currently still 

being analyzed but microsatellite 

amplification of Florida panther DNA 

was successfully extracted in 60% of the 

samples. Although collecting panther 

scat is labor intensive, utilizing DNA 

extracted from Florida panther scat holds 

promise as an unobtrusive technique to 

monitor the genetic health and individual 

makeup of the population. A final report 

will be completed later this year. 

Panther Peripheral Area Survey 

The only verified breeding population of 

Florida panthers is in the southern 

portion of the state, south of the 

Caloosahatchee River and Lake 

Okeechobee, in the Big Cypress and 

Everglades physiographic regions. This 

population has been growing since the 

mid 1990’s and so far 3 radio-collared 

panthers have crossed the river. 

Additionally, three uncollared panthers 

have been verified north of the river in 

recent years: two by tracks and/or 

photos, the other was road-killed. All of 

these have been males that dispersed 

from the core population. Only three are 

presumed to still be alive. Since resident 

male panthers typically encompass 

several females within their territory it is 

hypothesized that searching these ranges 

will afford the best opportunity of 

finding other panthers if they exist. This 

5-year study to determine the occurrence 

and status of panthers on peripheral 

areas of their presently known range has 

entered its final year. Systematic sign 

surveys have been conducted in areas 

where two male panthers had established 

territories. No sign of other panthers 


was found. A final report is scheduled 

for 2003. 

Effect of Genetic Introgression on 

Prevalence and Intensity of 

Gastrointestinal Helminth Infections in 

Florida Panthers 

The effects of genetic restoration of the 

Florida panther are being examined on 

many fronts. Complementing other 

projects, this study will indirectly assess 

the suspected improvement in immune 

function in intergrades by comparing 

gastrointestinal tract parasite burdens to 

that seen in original Florida panthers. 

Concurrently we will assess the efficacy 

of field anthelmintic treatment of 

panthers. Before introgression, 

gastrointestinal parasite burdens were 

assessed in 11original Florida panthers. 

Gastrointestinal tracts from a minimum 

of 7 panthers descended from Texas 

puma will be assessed by 2005 at which 

time a final report will be prepared. 

RECENT PUBLICATIONS 

CUNNINGHAM, M.W., M.R. DUNBAR, C.D. 

BUERGELT, B. HOMER, M. ROELKE- 

PARKER, S.K. TAYLOR, R. KING, S.B. 

CITINO, AND C. GLASS. 1999. Atrial 

septal defects in the Florida panther. 

Journal of Wildlife Diseases 35(3): 519- 

530. 

DEES, C.S., J.D. CLARK, AND F.T. VAN 

MANEN. 2001. Florida panther habitat 

use in response to prescribed fire. 

Journal of Wildlife Management 65:141- 

147. 

DUNBAR, M.R., M.W. CUNNINGHAM, AND 

S.T. LINDA. 1999. Vitamin A 

Concentrations in Serum and Liver from 

Florida Panthers. Journal of Wildlife 

Diseases 35(2): 171-177. 

JANIS, M.W. AND J.D. CLARK. 2002. 

Response of Florida panthers to 

recreational deer and hog hunting.

Journal of Wildlife Management. 

66:839-848. 

KRAMER, P.C. AND K.M. PORTIER. 2001. 

Modeling Florida panther movements in 

response to human attributes of the 

landscape and ecological settings. 

Ecological Modeling 140:51-80. 

LAND, E.D., D.R. GARMIN, AND G.A. HOLT. 

1998. Monitoring female Florida 

panthers via cellular telephone. Wildlife 

Society Bulletin. 26(1): 29-31. 

LAND, E.D. AND R.C. LACY. 2000. 

Introgression level achieved through 

Florida panther genetic restoration. 

Endangered Species Update 17: 99-103. 

MAEHR, D.S. 1998. The Florida panther in 

modern mythology. Natural Areas 

Journal. 18(2): 179-184. 

MAEHR, D.S. AND J.P. DEASON. 2002. 

Wide-ranging carnivores and 

development permits: constructing a 

multi-scale model to evaluate impacts on 

the Florida panther. Clean Technologies 

and Environmental Policy. 3:398-406. 

MAEHR, D.S., R.C. LACY, E.D. LAND, O.L. 

BASS, JR., AND T.S. HOCTOR. 2002. 

Evolution of population viability 

assessments for the Florida panther: a 

multiperspective approach. Pages 284- 

311 in: Population Viability Analysis. 

University of Chicago Press, Chicago. 

MAEHR, D.S., E.D. LAND, D.B. SHINDLE, 

O.L. BASS, AND T.S. HOCTOR. 2002. 

Florida panther dispersal and 

conservation. Biological Conservation 

106:187-197. 

MANSFIELD, K.G. AND E.D. LAND. 2002. 

Cryptorchidism in Florida panthers: 

prevalence, features, and effects of 

genetic restoration. Journal of Wildlife 

Diseases 38(4):693-698. 

ROTSTEIN, D.S., S. TAYLOR, J. HARVEY, J. 

BEAN. 1999. Hematologic effects of 

Cytauxzoonosis in Florida Panthers and 

Texas Cougars in Florida. Journal of 

Wildlife Diseases 35(3): 613-617. 



ROTSTEIN, D.S., R. THOMAS, K. HELMICK, 

S. CITINO, S. TAYLOR, M. DUNBAR. 

1999. Dermatophyte infections in freeranging 

Florida Panthers (Felis concolor 

coryi). Journal of Zoo and Wildlife 

Medicine 30(2): 281-284. 

ROTSTEIN, D.S., S.K. TAYLOR, J. BRADLEY, 

AND E.B. BREITSCHWERDT. 2000. 

Prevalence of Bartonella henselae 

antibody in Florida panthers. Journal of 

Wildlife Diseases 36(1):157-160. 

ROTSTEIN, D.S., S.K. TAYLOR, A. 

BIRKENHAUER, M. ROELKE-PARKER, 

AND B.L. HOMER. 2002. Retrospective 

study of proliferative papillary vulvitis 

in Florida panthers. Journal of Wildlife 

Diseases 38:115-123. 

TAYLOR, S.K., E.D. LAND, M. LOTZ, M. 

ROELKE-PARKER, S.B. CITINO, AND D. 

ROTSTEIN. 1998. Anesthesia of freeranging 

Florida panthers, 1981-1998. 

Proceedings of American Association of 

Zoo Veterinarians, Omaha, Nebraska. 

TAYLOR, S.K., C.D. BUERGELT, M.E. 

ROELKE-PARKER, B.L. HOMER, AND 

D.S. ROTSTEIN. 2002. Causes of 

mortality of free-ranging Florida 

panthers. Journal of Wildlife Diseases 

38:107-114. 


LAND, D., M. CUNNINGHAM, R. MCBRIDE, 

D. SHINDLE, AND M. LOTZ. 2002. 

Florida panther genetic restoration and 

management. Annual Report 2001- 

2002. Florida Fish and Wildlife 

Conservation Commission, Tallahassee. 

111pp. 

LAND, E.D. AND R.C. LACY. 2000. 

Introgression level achieved through 

Florida panther genetic restoration. 

Endangered Species Update 17: 99-103. 

MAEHR, D.S. 1990. The Florida panther 

and private lands. Conservation Biology 

4(2): 167-170. 

MAEHR, D.S., E.D. LAND, D.B. SHINDLE, 

O.L. BASS, AND T.S. HOCTOR. 2002.


Florida panther dispersal and 

conservation. Biological Conservation 

106:187-197. 

SEAL, U.S., AND WORKSHOP PARTICIPANTS. 

1992. Genetic management strategies 

and population viability of the Florida 

panther. Yulee, Florida: U.S. Fish and 

Wildlife Service. 

SEAL, U.S., editor. 1994. A plan for genetic 

restoration and management of the 

Florida panther (Felis concolor coryi). 

Conservation Breeding Specialist Group, 

Apple Valley, MN. 24pp. 


SHINDLE, D., D. LAND, M. CUNNINGHAM, 

AND M. LOTZ. 2001. Florida panther 

genetic restoration and management. 

Annual Report 2000-2001. Florida Fish 

and Wildlife Conservation Commission, 

Tallahassee. 102pp. 

TINSLEY, J.B. 1970. The Florida Panther. 

Great Outdoors Publishing Company. 

St. Petersburg, FL. 

YOUNG, S.P. AND E.A. GOLDMAN. 1946. 

The puma – mysterious American cat. 

Dover Publications, Inc., New York. 

385pp.

IDAHO MOUNTAIN LION STATUS REPORT 

STEVE NADEAU, Wildlife Staff Biologist, Idaho Department of Fish and Game, 600 South 

Walnut, Box 25, Boise, Idaho 83707, USA, email: snadeau@idfg.state.id.us 

INTRODUCTION 

Lions were classified as big game 

animals in 1972. The 1990 Mountain Lion 

Management Plan, called for the reduction 

in harvest of female lions, and maintain a 

harvest of approximately 250 lions 

statewide. However, lion harvest peaked 

statewide in 1998 when 798 lions were 

harvested. Consequently, a new lion plan 

was developed to address the changes in the 

populations and allow more hunting 

opportunity. Idaho completed the latest 

Mountain Lion Management Plan in 2002. 

The lion plan called for maintaining current 

lion distribution statewide as a goal. 

However, individual regions may adjust 

harvest to either increase or decrease 

populations depending upon the objectives 

for that area. Seasons were made more 

lenient, running from August 30 – March 31 

in most units. In some areas, 2-lion bag 

limits were initiated. Hounds were allowed 

in most units, and non-resident hound 

hunting was expanded. Female quotas were 

still used in most of the southern part of the 

state. 

HISTORY 

The legal status and public perception of 

mountain lions in Idaho has changed over 

time. In the late 1800’s and early 1900’s, 

mountain lions and other predators such as 

wolf, coyote, grizzly and black bears were 

perceived as significant threats to livestock 

and human interests and were systematically 

destroyed. Between 1915 and 1941, hunters 

employed cooperatively by the State, 

livestock associations, and the Federal 

25 


Government killed 251 mountain lions in 

Idaho; the take by private individuals is not 

known. During the period 1945-1958, 

bounties were paid for mountain lions in 

Idaho with an annual average of 80 

mountain lions turned in for payment 

(Figure 1). The 1953-54 winter period 

yielded the highest recorded bounty harvest 

of 144 mountain lions (Figure 1). Bounty 

payments ranged from $50 in the early 

1950’s to $25 per lion during the last 4 years 

of payments. 

Mountain lion sport harvest became 

increasingly popular after 1958. Average 

annual harvest was estimated at 142 lions 

from 1960 through 1971 (Figure 2). During 

this period there were no restrictions or 

regulations on the harvest of mountain lions. 

An estimated 303 lions were harvested 

during the 1971-72 season. 

Research conducted by Maurice 

Hornocker in the Frank Church River of No- 

Number of Lions Killed 

160 

140 

120 

100 

80 

60 

40 

20 

0 

1950 

1951 

1952 

1953 

1954 

1955 

1956 

1957 

1958 

1959 

Figure 1. Mountain lion bounty records, 

1950 – 1959. From 1950-1954 bounty was $50 

per lion; 1955-1959 the bounty was $25 per 

lion.

26 IDAHO MOUNTAIN LION STATUS REPORT · Nadeau 

Lion Harvest 

350 

300 

250 

200 

150 

100 

50 

0 

1960 

1962 

1964 

1966 

1968 

1970 

1972 

1974 

1976 

Unregulated Harvest Regulated 

1978 

1980 

Figure 2. Unregulated mountain lion harvest 

from 1960-71, and regulated harvest from 

1972 -1981. 

Return Wilderness from 1964-1973 added 

significantly to our knowledge. As a result 

of the research, the mountain lion was 

reclassified as a big game species in 1972. 

Harvest was then able to be regulated and 

resulted in some closed units, bag limits, and 

shortened seasons. Mandatory reporting 

was started in 1973, and a tag has been 

required since 1975. 

Populations of elk and deer continued to 

increase across the state during the 1980’s 

and early 1990’s, and the resulting mountain 

lion population did as well. The apparent 

increase in lion populations allowed the 

department to increase opportunity for 

harvest. Harvest continued to increase as a 

result of liberalized seasons and increased 

populations and peaked in 1997 (Figure 3). 


Lions were distributed across most of the 

suitable habitat in the state (Figure 4). 

Management tended to keep lion 

populations at a low density in developed 

areas or areas with high road density. 

However, most of the areas that received 

high harvest lay adjacent to lightly roaded 

reservoir areas that seemed to continue to 

provide dispersing animals. Distribution 

900 

800 

700 

600 

500 

400 

300 

200 

100 

0 

1982 

1984 

1986 

1988 

1990 


Harvest 

1992 

Year 

1994 

1996 

1998 

2000 

Lion Harvest 

Figure 3. Statewide mountain lion harvest. 

The year on the x-axis represents the date the 

season started, i.e. seasons run from fall 

through spring. 

Figure 4. Statewide mountain lion harvest by 

management unit and lion DAU where 

rankings are based on lions harvested/100mi 2 

where very low=. 03, low=. 3-.5, moderate=. 

6-1.0, high=1.1-2.0, and very high=2.6-3.0. 

The shaded units have female lion quotas.

appeared to be somewhat stable, though 

overall abundance may be declining. 

Population estimates have not been 

made for Idaho in recent years, though some 

radio collaring mortality information in 

Idaho indicated a high rate of sustainable 

harvest in some areas. Given an estimated 

harvest rate statewide of approximately 

15%, we would estimate approximately 

4,600 lions (+ 2,000). Research has been 

ongoing to attempt to develop a population 

index, however, nothing has been finalized 

(Zager et al. 2002). All lions harvested must 

be reported. Pelts were tagged and a 

premolar was removed for aging. Prior to 

2000, lion ages were estimated using tooth 

drop measurements. Based on various tests, 

tooth sectioning replaced tooth drop as a 

more reliable estimate of age and has been 

used since 2000. For data analysis purposes, 

units were grouped by similar characteristics 

into Data Analysis Units (DAUs). Age data 

and harvest rates were used to attempt to 

identify population trends for a lion by 

DAU. Populations modeling using these 

harvest data were used to estimate 

population demographics and relative 

abundance. 

Lion densities were highest in the 

northern part of the state where white-tailed 

deer and elk were common. Harvest by 

DAU size was used to standardize and 

compare lion harvest rates and estimated 

lion abundance (Figure 4). 


There were 99 big game management 

units in Idaho, which were grouped into 18 

mountain lion management DAUs (Figure 

4). The southern part of the state was 

predominantly managed under a female 

quota system, and the northern part of the 

state was mostly general hunts with most 

seasons running from August 30 – March 

31. Quotas and seasons were set by unit or 

DAU, usually based on historical harvest 

rates, big game objectives, depredations, 


IDAHO MOUNTAIN LION STATUS REPORT · Nadeau 27 

perceived lion population condition, lion 

hunter success rates and perceptions, public 

input, and commission desires. 

Biological objectives for lions were not 

well established by DAU, though age data 

were collected on all lions harvested. A 

minimum of 20% males 5+ years of age in 

the harvest was established as a test 

objective in some DAUs to adaptively 

manage populations by attempting to grow 

or reduce populations through harvest 

management, and monitor resultant age 

structures in the harvest. Regional wildlife 

managers in the state were given a great deal 

of flexibility to be able to set objectives for a 

given DAU. Lion harvest increased steadily 

through the 1980’s and 1990’s and peaked at 

798 mountain lions harvested in 1997. Lion 

harvest declined in most areas of the state 

following the 1997 season despite a 

liberalized lion hunting season in most of 

the state (Figure 3). 

Hunting with hounds accounted for 

about 80% of the annual lion harvest in 

Idaho. The rest of the harvest occurred 

incidentally to other big game hunting 

(13%), spot and stalk (5%), or predator 

calling (1%). The use of electronic calls 

was allowed in 2 management units where 

predation was a concern and access was 

limited. Dogs were prohibited through 

much of the general deer and elk rifle 

seasons. Pursuit with dogs was allowed in 

units with female quotas once the quota was 

reached. In a few of these units, hunting for 

males was allowed once the female quota 

was reached. 

Mountain lion tag sales increased 25% 

from 1998–2002, and in 2002 were at an all 

time high of 20,640 total tags sold (Table 1). 

Reduced prices, increased nonresident sales 

of special tags, and liberalized seasons and 

nonresident hound hunter regulations all 

added to increased sales. Additionally, in 

some parts of the state outfitters were 

engaged to increase harvest of lions to help

28 IDAHO MOUNTAIN LION STATUS REPORT · Nadeau 

Table 1. Mountain lion tag sales in Idaho 

from 1998 through 2002. 

Year Resident 

Tags 

Nonresident 

Tags 

Total 

Tags 

Sold 

1998 16,196 351 16,547 

1999 17,072 813 17,885 

2000 18,369 961 19,330 

2001 18,561 888 19,449 

2002 19,757 883 20,640 

reduce predation problems on elk and 

bighorn sheep. 


CONFLICTS 

Currently, Idaho law allows for killing 

lions or bears that are in the act of 

“molesting” livestock. This law also 

requires that lions killed in this fashion need 

to be reported to the Department. Idaho law 

also allows lions that are perceived as 

threats to human safety to be killed. 

Department policy provides that lions that 

have caused problems or have depredated 

should be captured and euthanized. Most 

depredations are reported to U.S. Wildlife 

Services and they handle the removal. 

Policy also provides that lions that present a 

threat due to proximity to residential 

housing or other area of human habituation 

or activity should be moved or chased in a 

preemptive fashion. Depending on the 

circumstance, if the animal has become 

habituated or caused problems, the lion can 


be destroyed. Orphaned kittens are not 

rehabilitated for release back into the wild. 

Idaho averaged 3-4 safety related 

complaints annually from 1998-2002 and 

about 50% required capture or removal of a 

lion. There has been 1-recorded human 

injury in Idaho caused by lions, and that 

occurred in 1999 to a 13-year-old boy. 

Lion related depredations that required 

compensation averaged about 1-2 per year. 

Average annual compensation form 1998- 

2002 was $4717 for lion depredations on 

livestock. During that same time, 46 lions 

were removed due to depredation situations. 

RESEARCH 

The Department has been researching 

techniques for population monitoring in 

north central Idaho by conducting aerial 

track surveys (Gratson and Zager 2000), and 

a mark-recapture technique using rub 

stations and biopsy darts (Zager et al. 2002). 

These efforts are still preliminary in nature. 


GRATSON, M.W., AND P. ZAGER. 2000. Elk 

ecology. Study IV. Factors influencing 

elk calf recruitment. Job No. 2. Calf 

mortality causes and rates. Federal Aid 

in Wildlife Restoration, Job Progress 

Report, W-160-R-26. Idaho Department 

of Fish and Game, Boise. 

ZAGER, P., M.W. GRATSON, AND C. WHITE. 

2002. Elk ecology. Study IV. Factors 

influencing elk calf recruitment. Job No. 

2. Calf mortality causes and rates. 

Federal Aid in Wildlife Restoration, Job 

Progress Report. W-160-R-29. Idaho 

Department of Fish and Game, Boise.

MONTANA MOUNTAIN LION STATUS REPORT 

RICH DeSIMONE, Montana Fish, Wildlife & Parks, 1420 East Sixth Avenue, Helena, MT 

59620, USA, email: rdesimone@state.mt.us 

ROSE JAFFE, Montana Fish, Wildlife & Parks, 1420 East Sixth Avenue, Helena, MT 59620, 

USA, email: rjaffe@state.mt.us 

INTRODUCTION 

Mountain lions in Montana are classified 

as a big game species. Overall management 

direction is provided in the Montana Fish, 

Wildlife & Parks’ (MFWP) 1996 

Environmental Impact Statement (EIS) – 

Management of Mountain Lions in 

Montana. According to the EIS, objectives 

concerning lion management are “… to 

maintain mountain lion and prey 

populations, to maintain mountain lion 

populations at levels that are compatible 

with outdoor recreational desires, and to 

minimize human-lion conflicts and livestock 

depredation”. 


Figure 1. Montana mountain lion hunting districts. 

29 


Mountain lions are currently distributed 

over approximately 75% of the state. Lions 

have filled habitats in western and central 

Montana and are continuing to expand in the 

eastern part of the state. Montana does not 

estimate lion populations, however, trends 

are monitored through harvest/mortality 

data, tooth age information, damage/conflict 

reports, and information from houndsmen. 


Lion harvest objectives are guided by 

balancing concern for human safety and 

demand for sport hunting. Montana’s 155 

deer and elk hunting districts are combined 

into 74 mountain lion hunting districts 

(Figure 1).

30 MONTANA MOUNTAIN LION STATUS REPORT · DeSimone and Jaffe 

Table 1. Montana lion hunting statistics, 1998-2002. 

1998 1999 2000 2001 2002 

License sales 

Resident 5421 5886 5138 5116 6337 

Non-resident 510 519 493 421 281 

Total 5931 6405 5631 5537 6618 

Lion Quota 

Harvest 

868 758 661 620 581 

Female 417 335 293 252 188 

Male 351 319 291 257 219 

Unknown 8 0 0 0 0 

Total 776 654 584 509 407 

Harvest is regulated through quotas and 

only one lion can be taken per hunter per 

year. Quotas include any lion, male and 

female, and female sub quotas. During the 

fall hunting season (last week of Oct 

through Nov), the use of dogs is not 

allowed. Harvest during the fall season has 

been in affect for 4 years and less than 10 

lions were harvested each year (Table 1). 

Hunting with dogs is allowed during the 

winter season (Dec 1 – Apr 14) and accounts 

for over 95% of the harvest. Licensed 

hunters are also allowed to chase lions 

during the winter season. Recent legislation 

will allow the purchase of non-harvest chase 

licenses. 


INTERACTION/CONFLICTS 

MFWP’s Mountain Lion Depredation 

Table 2. Montana mountain lion incidents and removals, 1998-2002. 


and Control Guidelines are used to deal with 

different types of incidents. Depending on 

the situation, management actions include 

education, relocation, and removal (Table 

2). Montana does not pay for losses 

attributed to lions. 

RESEARCH 

Garnet Mountains – Mountain Lion 

Research, 1998 – present. 

The goal is to document the influence of 

hunting on population characteristics and 

evaluate the ability of various survey 

techniques to detect trends in lion 

abundance. 

1998 1999 2000 2001 2002 

Incidents 1 

Public safety 41 18 37 30 20 

Depredation 2 58 44 35 37 29 

Total 

Removals 

99 62 72 67 49 

Public safety 20 2 3 5 2 

Depredation 30 20 20 11 14 

Total 50 22 23 16 16 

1 

Incident: A conflict between a human and lion that may have serious results (i.e. a lion killing a dog or a lion that must be 

forced to back down). 

2 

Depredation: Includes death of pets and death and injury of livestock.

NEVADA MOUNTAIN LION STATUS REPORT 

RUSSELL WOOLSTENHULME, Nevada Department of Conservation & Natural Resources, 

1100 Valley View Road, Reno, NV 89512, USA, email: rwoolstenhulme@ndow.org 

INTRODUCTION 

The Nevada Division of Wildlife 

completed its Comprehensive Mountain 

Lion Management Plan in January 1995. 

The Nevada Board of Wildlife 

Commissioners approved the plan in 

October of that year. The plan is scheduled 

for revision during 2003. 

The goals and objectives of the mountain 

lion plan are to maintain lion distribution in 

reasonable densities throughout Nevada, to 

control mountain lions creating a public 

safety hazard or causing property damage, 

and to provide recreational, educational and 

scientific use opportunities of the mountain 

lion resource. Additional goals include 

maintaining a balance between mountain 

lions and their prey, and finally to manage 

mountain lions as a metapopulation. 

The mountain lion’s legal classification 

in Nevada was changed by regulation from 

unprotected (predator) to game animal in 

1965. The change in classification resulted 

in the requirement of a valid hunting license 

to hunt mountain lion, along with some 

restrictions in the method of take. This 

provision precluded the taking of lions at 

any time other than from sunrise to sunset 

and it also defined legal weapons as 

shotgun, rifle, or bow and arrow. The 

season was defined as either sex, yearround, 

and no limit was set nor was a tag 

required. Mountain lion harvest 

management has changed substantially from 

1965 to the present. 

In 1968, a tag requirement was 

instituted, and although no limits were 

established, it became possible to record 

31 


sport hunter harvest. Another major change 

occurred in 1970, when a limit of one lion 

per person was set, and a six-month season 

was established. During that year, the 

requirement that all harvested lions be 

validated by a representative of the 

Department within five days after the kill 

was also established. This regulation 

presented the Department the first real 

opportunity to collect biological data from 

the mountain lion. 

In 1972, the Nevada Department of 

Wildlife initiated a study of the mountain 

lion as a part of the Ruby-Butte deer project 

in eastern Nevada. The objective was to 

determine the status of lion populations 

within this high-density deer area, and, to 

evaluate them in relation to deer 

populations. Within two years, this 

objective was changed to: a) establish 

population estimates of mountain lions by 

mountain range or management area 

statewide, b) establish basic habitat 

requirements, c) establish a harvest 

management program. From that period on, 

increased emphasis was placed upon lion 

capture and marking with the more 

sophisticated telemetry devices which were 

being manufactured. This program involved 

lion monitoring from both land and air and 

was instrumental in expanding our life 

history information base, as well as 

providing an approach toward estimating the 

annual population status in key mountain 

ranges. The findings from this study were 

then utilized in formulating an approach 

toward estimating statewide lion 

populations. This ten (10) year study formed

32 NEVADA MOUNTAIN LION STATUS REPORT · Woolstenhulme 

the basics for most management activities 

that have been implemented since 

publication of this study in 1983. 

In 1976, 26 mountain lion management 

areas were described statewide, and a 

harvest quota established for each to control 

the sport harvest. This “Controlled Quota 

Hunt” was the most restrictive season ever 

established for mountain lion in Nevada. 

In 1979, the “Controlled Quota Hunt” 

was modified utilizing six management 

areas whereby a harvest objective was 

established which allowed the hunting of 

lions in each of the six areas until the 

predetermined number of lion were taken. 

In 1981, the “Harvest Objective” hunting 

season concept was applied statewide. 

Initially this system required a hunter to 

obtain a free hunt permit for the opportunity 

to hunt in one (1) management area. In 

1994, hunters were allowed to obtain a free 

hunt permit that authorized the hunter to 

hunt in two (2) management areas until the 

established harvest objective was reached. 

Both of these permit systems allowed 

hunters to change management areas at will 

as long as the harvest objective had not been 

reached in the desired management area(s). 

In 1995, the hunt permit approach was 

modified to eliminate the physical issuance 

of a permit in favor of establishing a 1-800 

telephone number. This system allows 

hunters to hunt in any management area in 

which the harvest objective has not been 

reached. The hunter must, however, call the 

1-800 number before starting to hunt to 

determine which management area(s) are 

still open to hunting. 

In 1997, changes were made to mountain 

lion regulations to increase mountain lion 

harvest, while maintaining the integrity of 

the harvest objective limits system. Those 

changes included the reduction of tag fees, 

over-the-counter tag sales, increasing bag 

limits from one tag per hunter to two tags 

per hunter, and consolidation of some of the 


harvest unit groups. 

In 1998, Nevada’s southern region was 

modified to provide for a year-round hunting 

season on mountain lions. The entire state 

went to a year-round season in 2001. 

New changes were made again for the 

2003 season. These changes modified 

harvest unit groups from 24 groups 

throughout the state to three statewide 

regions corresponding with the Division’s 

three management regions. The mountain 

lion season continues to be year-round but 

season dates were changed to March 1 st of 

each year to the last day of February, 

corresponding with the dates on a Nevada 

hunting license. 


Mountain lions seem well adapted to the 

wide variety of habitat and environmental 

conditions that exist in Nevada. They have 

been observed to live or wander through 

almost every mountain range from the 

Mojave Desert in extreme southern Nevada 

to alpine forests at the highest elevations in 

the northern part of the state. Distribution 

appears to be primarily influenced by prey 

availability, and has remained fairly 

consistent through time. 

Mountain lion populations are estimated 

utilizing a life table model (retrospective 

harvest/ mortality). The model utilizes 

known harvest/ mortality rates and 

recruitment rates (as determined from markrecapture 

and telemetry studies) to calculate 

a retrospective estimate of minimum viable 

population size needed to sustain known 

harvest rates over the same time period. 

Although no defined confidence limit is 

used during this process, our confidence in 

this model is relatively high based on the 

fact that harvest rates have continued over 

time at a constant rate without signs of 

extirpation, reduced harvest rates, or 

increased average age of harvested lion. 

Based on our current estimation methods,

NEVADA MOUNTAIN LION STATUS REPORT · Woolstenhulme 33 

Table 1. Mountain lion tag sales, sport hunter harvest, and hunter success by class of hunter. 

Tag Sales Harvest Hunter Success 

Year Resident Nonresident Total Resident Nonresident Total Resident Nonresident Total 

1998 643 124 767 73 67 140 11% 54% 18% 

1999 680 109 789 70 56 126 10% 51% 16% 

2000 883 169 1052 104 81 185 12% 48% 18% 

2001 838 98 936 103 58 161 12% 59% 17% 

2002 1030 202 1232 105 63 168 10% 31% 14% 

2003 1060 131 1191 89 39 128 8% 30% 11% 

Total 5,134 833 5,967 544 364 908 11% 44% 15% 

Average 856 139 995 91 61 151 11% 44% 15% 

lion populations within Nevada are between 

3000-4000 animals. 


Mountain lions have been classified as a 

big game species since 1965. They have 

been hunted annually since that time. A 

Nevada resident mountain lion tag costs 

$25.00, and a Nevada nonresident mountain 

lion tag costs $100.00. On the average, 

nonresident hunters account for 

approximately 14% of tag sales, but harvest 

a greater proportion of lions than do resident 

hunters (Table 1). Total sport hunter harvest 

has averaged 151 lions per year for the last 6 

years (Table 2). 

The open season for hunting mountain 

lions in Nevada currently runs year-round 

(March 1 – last day of February) (Table 3). 

Table 2. Mountain lion harvest by harvest type and sex. 


Any legal weapon may be used to harvest a 

mountain lion, and dogs may be used to hunt 

a mountain lion under the authority of a 

current State of Nevada hunting license and 

mountain lion tag. Because the mountain 

lion season is year-round no pursuit only 

season exists. A resident or a non-resident is 

eligible to obtain two mountain lion tags 

each year. A person who harvests a 

mountain lion in Nevada must, within 72 

hours after harvesting it, personally present 

the skull and hide to a representative of the 

division for inspection. The representative 

shall affix a State of Nevada mountain lion 

seal permanently to the hide. A seal must be 

permanently affixed to the hide of a 

mountain lion before it can be possessed by 

an individual or removed from the state. It 

is unlawful to kill a female mountain lion 

Sport Hunter Harvest Depredation Take Total 

Year Male Female Total Male Female Total Male Female Total 

1998 85 55 140 12 8 20 97 63 160 

1999 77 49 126 12 10 22 89 59 148 

2000 102 83 185 8 3 11 110 86 196 

2001 95 66 161 8 8 16 103 74 177 

2002 99 69 168 10 16 26 109 85 194 

2003 77 51 128 7 8 15 84 59 143 

Average 89 62 151 10 9 18 99 71 170


Table 3. Nevada Mountain Lion Units and Quotas 2003 – 2005. 

Unit Group 

UNIT 1 (Western Region) 

011 - 015, 021, 022, 031, 

032, 034, 035, 041 - 046, 

051, 181 – 184, 192, 194 - 

196, 201 - 206, 291 

2003-2004 Season 

Dates 

March 1, 2003 – 

Feb 29, 2004 

2003-2004 

Harvest Objectives 

114 

2004-2005 Season 

Dates 

March 1, 2004 – 

Feb 28, 2005 


2004-2005 

Harvest Objectives 

033 Closed 0 Closed 0 

UNIT 2 (Eastern Region) 

061, 062, 064 – 068, 071 - 

078, 081, 101 – 108, 111 – 

115, 121, 131 – 134, 141 – 

145, 151, 152, 154, 155 

079* 

UNIT 3 (Southern Region) 

161 - 164, 171 - 173, 211, 

212, 221 – 223, 231, 241 – 

244, 251 - 253, 261 - 268, 

271 – 272 

March 1, 2003 – 

Feb 29, 2004 

March 1, 2003 – 

Feb 29, 2004 

March 1, 2003 – 

Feb 29, 2004 

163 

4 

68 

March 1, 2004 – 

Feb 28, 2005 

March 1, 2004 – 

Feb 28, 2005 

March 1, 2004 – 

Feb 28, 2005 

280 – 284 Closed 0 Closed 0 

* Interstate hunt with Utah. Nevada and Utah hunters may hunt within open units in both states. Nevada hunters 

hunting in Utah must abide by Utah regulations. 

that is accompanied by a spotted kitten, or to 

kill or possess a spotted mountain lion 

kitten. It is also unlawful in Nevada to trap 

a mountain lion, if a mountain lion is 

accidentally trapped or killed, the person 

trapping or killing it shall report the trapping 

or killing within 48 hours to the division. 

The animal must be disposed of in 

accordance with state law. 

Mountain lion harvest objectives are 

calculated for each administrative region on 

a semi-annual basis using standardized 

methodology. Harvest objectives are 

calculated and recommended in order to 

achieve a specific management action over a 

short-term period (no more than two years). 

Management actions may be designed to 

increase, stabilize and maintain, or decrease 

mountain lion populations within each of the 

three administrative regions in Nevada. 

114 

163 

Calculations of harvest objectives by 

administrative region incorporate the use of 

scientific data to determine the current 

population trend and population density. A 

“political index” may be employed to adjust 

harvest objectives within smaller geographic 

areas (big game management areas) in order 

to achieve the desired management goal. 

Biologists make annual adjustments to 

harvest objective recommendations for each 

administrative region only after careful 

review of the following data and information 

that is collected, assembled and distributed 

by the Game Bureau by October of each 

year. 

A. Data used to assess population trend, 

including, but not limited to: 

4 

68 

1) The current regional population 

model.

2) Sex, weight and age data from 

harvested mountain lions for the 

previous recording period (March 1 

- February 28). 

B. Data used to assess population density, 

including, but not limited to: 

1) The current regional population 

model. 

2) Data showing the unit of effort to 

observe or harvest mountain lions. 

3) Average weight information, 

comparing weights of harvested 

animals by sex and cohort group to 

the long-term data set (1968 - 

2003). 

C. Data to quantify “bio-political” 

considerations, including, but not 

limited to: 

1) A summary of the public safety 

complaint forms involving 

mountain lions as received by the 

Bureau for the previous recording 

period. 

2) A report of damage to private 

property caused by mountain lions 

as annually prepared by ADC. 

3) A prey species accounting 

spreadsheet as prepared by the 

region for the previous recording 

period. Adjustments from the 

baseline harvest objective level for 

each administrative region will be 

recommended in order to achieve 

the short-term (two-year) goal of 

maintaining, increasing, or 

decreasing mountain lion 

populations within the respective 

administrative region, utilizing 

harvest management as the primary 

tool to achieve the desired 

population goal. 

See Figure 1 for State of Nevada 

mountain lion hunt unit reference map. 


Figure 1. Nevada mountain lion hunt unit 

reference map. 




The Nevada Division of Wildlife 

Comprehensive Management Plan 

specifically addresses policy and procedure 

for dealing with nuisance or problem 

mountain lions. 

The Division of Wildlife is responsible 

by statute for controlling wildlife causing 

damage to personal property or endangering 

personal safety. The Division also has a 

responsibility to provide sport-hunting 

opportunities to Nevada sportsmen. This 

protocol sets forth procedures to be followed 

in controlling and preventing lion damage, 

addressing public safety issues and 

responding to sport hunting opportunities. 

In carrying out this policy where mountain 

lion/human interactions are involved, agents 

shall have the discretion to choose the most


applicable management option, following 

guidelines outlined in this protocol. All 

efforts will be directed at the individual lion 

causing the problem. 

Mountain lion/human interactions have 

increased throughout the West and in 

Nevada in the last several decades. During 

the same period, the number of depredation 

complaints and the number of lions taken on 

depredation complaints has also increased. 

The Division desires to reduce multiple 

depredations from the same animal and 

prevent harm to humans. 

The Division recognizes three distinct 

categories of mountain lions involved in 

human/lion interactions. 

A. Nuisance Lion - a lion involved in a 

direct meeting with a human but did 

not exhibit aggressive behavior toward 

the human, a lion repeatedly observed 

in an area, or a situation where 

personal property is at risk. 

B. Depredating Lion - a lion that has 

injured or killed livestock or domestic 

pets. 

C. Dangerous or Aggressive Lion - a 

lion that has exhibited aggressive 

behavior towards humans. A lion that 

has an unnatural interest in humans 

without provocation and is perceived 

to be a threat to public safety. A lion 

located in a place or situation where 

human safety is of concern may be 

considered dangerous. 

Various management options are 

available to Division employees when a 

mountain lion conflict arises. The Division 

employee responding to or assigned to 

handle a lion/human conflict will have the 

primary responsibility to assess mountain 

lion involvement in an incident and conduct 

the necessary investigation. Agents may be 

required to make an assessment "on the 

spot" or if time permits make an assessment 


with consultation. 

At all opportunities, the Division will 

provide educational and informational 

materials to individuals concerned with lion 

management and people-lion conflict 

prevention. These materials will include 

options for pet and livestock protection and 

avoidance of dangerous encounters with 

mountain lions. Site-specific education and 

prevention efforts will be made in historic 

conflict areas. 

A field response by either a Division 

employee or his/her designated agent is 

required for all lion/human interactions 

involving the categories of lions defined. 

1. Nuisance Mountain Lions 

a. No management action combined 

with education effort. 

b. Deterrent methods combined with 

education effort. 

c. Capture, mark and relocate cougars 

if deterrent methods are 

unsuccessful or impractical. Lions 

identified for relocation will be 

transported to the following release 

sites in priority order. 

1) Instate release locations within 

low conflict areas 

2) Out of state governmental 

agencies 

3) University or research facilities 

4) Zoological gardens or Zoos 

d. Nuisance lions will be destroyed if 

relocating or deterrent methods are 

unsuccessful or impractical. 

2. Depredation Mountain Lions 

a. No management action combined 

with education effort. 

b. Deterrent methods including 

prevention materials (if applicable) 

combined with education effort. 

c. Capture, mark and relocate cougars 

if deterrent methods are 

unsuccessful or impractical. Lions

identified for relocation will be 

transported to the following release 

sites in priority order. 




agencies 



d. Depredating lions will be destroyed 

if deterrent methods or live capture 

is unsuccessful or impractical. 

3. Aggressive (Dangerous) Mountain 

Lions 

a. If a lion is dangerous because of its 

location and not its behavior it may 

be trapped, marked and relocated. 

If a lion is frequenting a city or 

town, it may be destroyed if capture 

methods fail or are impractical. 

Lions identified for relocation will 

be transported to the following 

release sites in priority order. 




agencies 



b. If the mountain lion is dangerous 

because it has exhibited aggressive 

behavior toward humans or is 

otherwise perceived to be a threat to 

human safety, or if the lion is 

involved in an attack on a human, 

destroy and necropsy the lion. 

Lions exhibiting aggressive 

behavior in remote areas should not 

be killed but instead an aggressive 

publicity and educational campaign 

should be made to alert people of 

the danger in the remote area and 

promote human avoidance of the 

area over the short-term. 



c. A detailed narrative report on each 

incident involving handling of 

dangerous lions will be prepared by 

the agent in control of the incident 

and forwarded to the Supervising 

Regional Game Biologist. 

Mountain lion incidents involving 

attacks or injury to people will be 

immediately reported through the 

chain of command to the Regional 

Manager, Administrator, Chief of 

Game and Chief Game Warden. 

All lions destroyed will be reported 

on the 351-harvest form. A copy of 

the detailed report, including any 

necropsy, coroner's report or other 

supporting information shall be sent 

to the Game bureau staff biologist 

responsible for mountain lions. A 

lion/human interaction form will be 

completed for each interaction. 

In those incidences where control 

becomes necessary, a regional list of persons 

who have requested consideration and are 

qualified to do control work, including 

private hunters/ trappers and outfitters/ 

guides will be a source of control, as well as 

U.S.D.A. APHIS/ADC personnel. 

Hunters/trappers, outfitters/guides or 

U.S.D.A. agents will not initiate control 

unless requested to do so by the Division. 

Hunters or trappers may be authorized to 

control problem animals during open or 

closed seasons. The hunter or trapper will 

buy a license and tag for use during the open 

season until the hunter or trapper's tag is 

filled. The hunter may continue control 

work after the tag is filled only under the 

authority of a depredation permit. Hunting 

during a closed season will be conducted 

only under the authority of a depredation 

permit. Depredation permits will only be 

issued to landowners/livestock owners for 

the control of specific depredating lions. 

Hunters or trappers may keep the lion if 

harvested under the authority of a valid


license and tag. All other lions become the 

property of the State. 

The USDA, APHIS/ADC, may be 

contacted to do control work any time of 

year. The APHIS/ADC agent shall attempt 

to control only the animal(s) causing 

damage. The agent will use discretion in the 

control of young animals. All lions taken by 

APHIS/ADC are the property of the State. 

A mountain lion harvest report form is 

completed for all mountain lion mortalities. 

A mountain lion/human interaction form is 

completed for all lion/human interactions. 

Records of lion mortality and human/ lion 

interactions are kept in computer databases 

in Reno. 


ERNEST, HOLLY B., WALTER M. BOYCE, 

VERNON C. BLEICH, BERNIE MAY, SAN 

J. STIVER, AND STEVEN G. TORRES. In 

Press. Genetic structure of mountain 

lion (Puma concolor) populations in 

California. Journal of Conservation 

Genetics. 


This paper used 412 samples from 

California and 19 samples collected in 

western Nevada within 50 km of California. 

The work helped define the geographic 

ranges of mountain lion populations in 

California. Population structure differed 

greatly by region - mountain lions in many 

California regions have significant barriers 

to genetic interchange and therefore are very 

different from one population to another. 

This paper, plus the work done for the 

Nevada DOW report indicate that 

populations in Nevada tend not to have as 

much obstruction to genetic interchange as 

those in most ecological regions of 

California, in general. This study shows that 

mountain lion management and conservation 

efforts should be individualized according to 

region and incorporate landscape-level 

considerations to protect habitat 

connectivity. 

A follow up study by Dr. Holly Ernest, 

on mountain lion genetic variation and 

phylogeography in Nevada is currently 

being finalized for future publication.

NEW MEXICO MOUNTAIN LION STATUS REPORT 

RICK WINSLOW, Large Carnivore Biologist, New Mexico Department of Game and Fish, P.O. 

Box 25112, Santa Fe, New Mexico 87504, USA, email: Rwinslow@state.nm.us 

INTRODUCTION 

The New Mexico Department of Game 

and Fish has almost completed 

implementation of its cougar management 

plan. We are beginning to develop a new 

five-year plan. 

Mountain lions have been classified as 

protected big game animals in New Mexico 

since 1971 and are currently hunted 

throughout most of the occupied habitat in 


Mountain lion management in New 

Mexico is multi-faceted. The department is 

attempting to develop a conservation 

strategy that allows both hunting and 

enjoyment of cougars by the non-hunting 

public. We also need to balance differing 

management issues: (1) depredation control 

to minimize economic losses to livestock 

operators, (2) minimizing human/cougar 

conflicts, (3) cougar removal where 

increasing deer and bighorn populations is 

the management priority. 

In 1999 we initiated a zone quota system 

for harvest management. Our zone quotas 

are based upon management decisions for 

either increasing, maintaining stable, or 

decreasing lion populations. 


Since their protection as a big game 

animal in 1971, mountain lions have steadily 

returned to suitable habitat throughout the 

state. Mountain lions generally inhabit the 

rougher country in New Mexico avoiding 

the low elevation desert areas and eastern 

plains. They do however occur in these 

areas in conjunction with pockets of mule 

39 


deer and areas of topographic diversity. Our 

current estimate of the cougar population in 

New Mexico is approximately 2150 cougars 

derived by multiplying density estimates by 

(Logan et al. 1996) the estimated amount of 

mule deer habitat. For regional estimates, 

we use a population model based on rates of 

recruitment and mortality from Logan et al. 

(1996), quantity of habitat and population 

density. 

Since 1979, successful hunters have 

been required to present their cougar to a 

Department official within 5 days of their 

harvest to have the pelt tagged, a tooth 

collected for aging, sex verified, and other 

information gathered. Reports of cougar 

depredation and damage are also kept. 

Harvest strategies have varied during the 

32 years cougars have been classified as a 

game animal. In 1971 only the southwestern 

corner of New Mexico was open to cougar 

hunting with a bag limit of one cougar and a 

4-month season. More areas of the state 

were opened to cougar hunting and seasons 

lengthened in subsequent years. From 1979 

to 1983 the season was 11 months long 

statewide with a bag limit of 2 cougars. 

In 1983 a bill was introduced to the New 

Mexico House of Representatives to return 

the cougar to its status prior to 1971 as a 

varmint. It was tabled but the legislature 

requested that the department gather more 

information on the cougar’s status. Evans 

(1983) investigated harvest trends and 

population estimates and determined that 

cougar populations had probably declined. 

His determination and public opinion

40 NEW MEXICO MOUNTAIN LION STATUS REPORT · Winslow 

resulted in more conservative harvest 

strategies. 

In 1984 the cougar season was shortened 

to 3 months in most of the state, with longer 

seasons in units that had high numbers of 

depredation complaints. From 1985 until 

1999 the season was 4 months long 

throughout the state with a bag limit of one 

cougar. In 1999 the state instituted a zone 

management system with harvest objectives 

and quotas. The season was extended to 6 

months throughout the majority of the state 

with low-elevation bighorn sheep ranges 

open year round. 

In 2002, the cougar season remained at 6 

months with a 1 cougar bag limit throughout 

most of the state with the following 

exceptions: year around hunting in selected 

desert bighorn sheep areas and Rocky 

Mountain bighorn sheep areas in the 

southern part of the state and on private 

property, and year round hunting in specific 

units in the southeastern corner of the state 

that have historically suffered high 

depredation losses. The bag limit was 

increased to 2 cougars in bighorn areas. 

Currently the state has 15 cougar 


Figure 1. Cougar Harvest Management 

Zones in New Mexico during 2002-03. 

management zones (Figure 1). Each is 

managed through a quota system for 

increasing, decreasing or stable populations 

of cougar (Table 1). The ratio of males to 

females harvested generally equals 60:40 

Table 1. Cougar harvest objectives by management zone in New Mexico, 2002-03. 

Zone Game Management Units Included in Zone Harvest Objective 

A 2 and 7 14 

B 5 and 50-51 20 

C 43-46, 48-49, and 53-55 38 

D 41-42, 47, and 56-58 16 

E 9 and 10 16 

F 6 and 8 16 

G 13-14, and 17 17 

H 19, 20, and 28-29 3 

I 18, 30, 34, and 36-38 20 

J 15-16, 21, and 25 38 

K 22-24 22 

L 26-27 Unlimited 

M 31-33, and 39-40 5 

N 4 and 52 3 

O 12 3 

231 a 

a Not including unlimited areas.

(Table 2). Hunters tend to selectively 

harvest the larger male lions. 

Since sport hunting was implemented in 

1971, use of hounds has been allowed but 

cubs and females with cubs cannot be taken. 

At least 90% of the harvest is through hound 

hunting. There is no pursuit season. 

Approximately 2000 lion licenses are 

sold per year currently. This number has 

gone up during the past decade. There is no 

limit to the number of lion licenses sold per 

year. 

Since 1998 cougar depredation 

complaints have ranged from 28 to 45 per 

year with 1 to 20 cougars killed per year. 

Mountain lion depredation incidents are 

typically dealt with on a case-by-case basis 

NEW MEXICO MOUNTAIN LION STATUS REPORT · Winslow 41 

Table 2. Permits issued and cougars harvested in New Mexico, 1981-2003. 

Hunt Year 

Permits 

Issued 

Male Harvest 


in New Mexico. Department policy is to 

resolve depredation and to minimize 

property damage, conflict and threat to 

human safety. When department or Wildlife 

Services investigation confirms a 

depredation incident, a depredation permit 

may be issued. Generally, either snares or 

hounds are used to capture the offending 

animal. Lions involved in depredation 

incidents are destroyed. Landowners may 

also kill lions in defense of human safety or 

property. The southeastern corner of the 

state has a preventative control program, 

which is in effect in Unit 30 to reduce 

depredation on domestic sheep. The 

preventative control program destroyed 110 

mountain lions between 1989 and 1999 and 

Female 

Harvest 

Unknown Total Harvest 

1981-82 360 78 44 3 125 

1982-83 481 55 44 1 101 

1983-84 661 67 65 0 132 

1984-85 443 47 32 0 79 

1985-86 472 56 48 0 104 

1986-87 437 55 46 0 101 

1987-88 456 43 35 0 78 

1988-89 450 58 33 0 91 

1989-90 482 71 41 0 112 

1990-91 781 73 35 0 108 

1991-92 765 77 42 0 119 

1992-93 826 68 37 0 105 

1993-94 926 75 52 0 127 

1994-95 1145 87 61 2 150 

1995-96 842 74 45 0 119 

1996-97 980 114 62 1 177 

1997-98 974 108 58 2 168 

1998-99 1485 95 58 0 153 

1999-00 1702 98 58 0 156 

2000-01 NA 1 140 96 0 236 

2001-02 NA 1 127 91 1 219 

2002-03 NA 1 161 120 3 284 2 

1 

Not yet determined. 

2 

Numbers may not be complete.

42 NEW MEXICO MOUNTAIN LION STATUS REPORT · Winslow 

has continued in the years since. 

In situations where depredation cannot 

be confirmed, the district wildlife officer 

will offer advice and suggestions as to how 

the complainant can avoid incidents with 

lions. Lions captured for reasons other than 

depredation are relocated to another area of 


Human safety incidents with lions are 

rare in New Mexico. Any lion involved in a 

human safety type of incident would be 

destroyed if caught. 


Ligon (1926) conducted the first 

investigation on cougars in New Mexico and 

determined that they were uncommon but 

preyed heavily upon domestic animals and 

deer. Hibben (1937) investigated lion 

biology in northern and western portions of 

the state. Prey use and movements in the 

southwestern corner of the state were 

documented via radio telemetry in the 

1970’s (Donaldson 1975, Johnson 1982). 

Cougar ecology in Carlsbad Caverns 

National Park, New Mexico and the 

Guadalupe Mountains National Park across 

the border in Texas was studied from 1982- 

85 (Smith et al. 1986). Ecology and 

population dynamics of cougars in the San 

Andres Mountains of south central New 

Mexico were studied from 1985-95 (Logan 

et al. 1996). This was the most intensive 

investigation of desert-dwelling cougars 

ever conducted. Beausoleil (2001) reviewed 

historic and current status of mountain lions 

in New Mexico. 


BEAUSOLEIL, R.A. 2001. Status of the 

Mountain Lion in New Mexico, 1997- 

2000. New Mexico Naturalist’s Notes 

3(1) pp. 33-47. 

DONALDSON, B. 1975. Mountain lion 

research. Final Report, Pittman 

Robertson Project W-93-17, Work plan 

15, Job 1. New Mexico Department of 


Game and Fish, Santa Fe, New Mexico 

USA. 

EVANS, W. 1983. The cougar in New 

Mexico: biology, status, depredation of 

livestock, and management 

recommendations. Response to House 

Memorial 42. New Mexico Department 

of Game and Fish, Santa Fe, New 

Mexico USA. 

HIBBEN, F.C. 1937. A preliminary study of 

the mountain lion (Felis oregonenis 

spp.). University of New Mexico 

Bulletin, Biological Series 5(3) 5-59. 

JOHNSON, J. 1982. Mountain lion research. 

Final Report, Pittman Robertson Project 

W-124-R-4, Job 1. New Mexico 

Department of Game and Fish, Santa Fe, 

New Mexico USA. 

LIGON, J.S. 1927. Wild Life of New Mexico, 

Its Conservation and Management. 

Being a Report on the Game Survey of 

the State, 1926 and 1927. State Game 

Commission Department of Game and 

Fish, Santa Fe, New Mexico. 

LOGAN, K.A., L.L. SWEANOR, T.K. RUTH, 

AND M.G. HORNOCKER. 1996. Cougars 

of the San Andres Mountains, New 

Mexico. Federal Aid in Wildlife 

Restoration, Project W-128-R, for New 

Mexico Department of Game and Fish, 

Santa Fe, New Mexico USA. 

NEW MEXICO DEPARTMENT OF GAME AND 

FISH. 1997. Long range plan for the 

management of cougar in New Mexico. 

Federal Aid in Wildlife Restoration 

Grant W-93-R-39, Project 1, Job 5. New 

Mexico Department of Game and Fish, 

Santa Fe, New Mexico USA. 

SMITH, T.E., R.R. DUKE, M.J. KUTILEK, AND 

H.T. HARVEY. 1986. Mountain lions 

(Felis Concolor) in the vicinity of 

Carlsbad Caverns, New Mexico and 

Guadalupe Mountains National Park, 

Texas. Harvey and Stanley Associates 

Incorporated, Alvisa, Texas USA.

STATE OF SOUTH DAKOTA MOUNTAIN LION STATUS REPORT 

MIKE KINTIGH, Regional Supervisor, South Dakota Game, Fish & Parks, 3305 West South St., 

Rapid City, SD 57702, USA, email: Mike.Kintigh@state.sd.us 

INTRODUCTION 

South Dakota (SD) is currently 

developing a management plan for mountain 

lions. The second draft of this document is 

currently under review by South Dakota 

Department of Game, Fish and Parks (SD 

GFP) staff. This document is also available 

to the public and interested parties for 

review and comment. A copy can be 

obtained by contacting Regional Supervisor 

Mike Kintigh (listed above) or Dr. Larry 

Gigliotti at 605-773-4231. This summer/fall 

SD GFP will be taking further steps to 

solicit public comments on the management 

plan. 

Mountain Lions are currently classified 

as a State Threatened Species in SD. 

However, in July that classification will 

likely change significantly. Legislative 

action in January of 2003 closed a legal 

loophole by defining the lion as a big game 

animal. This action will facilitate the 

removal of the lion from the State’s 

Threatened Species List. Game 

Commission action is still required to 

finalize the delisting and this is expected to 

occur in early June. Forty-five days is 

required for any commission finalization 

actions to take effect and this will occur in 

mid July, after the legislative action 

becomes law on July 1, 2003. 

It is important to note that while the lion 

will be taken off the threatened species list 

in South Dakota, it will actually gain 

additional protection under law by being 

defined as a big game animal. Criminal 

penalties will increase from class 2 

misdemeanors to class 1 misdemeanors, 

43 


carrying higher fines and longer jail 

sentences. 

A misconception exists in that by 

classifying the lion as a big game animal a 

hunting season will immediately be 

implemented. This is absolutely false! The 

lion will continue to be fully protected as a 

big game animal with a continuously closed 

season until at some undetermined point 

when additional management decisions are 

made. 

South Dakota has many objectives 

concerning mountain lion management. 

They are as follows: 

A) Evaluate the Legal status of the 

Mountain Lion in SD by April 1, 

2003. 

B) Evaluate strategies for monitoring & 

censusing Mountain Lion 

populations in SD by 2005. 

C) Maintain a statewide database of Mt. 

Lion activity including sightings, 

human interactions, depredation 

events and lion mortality. 

D) Develop a list of Mt. Lion research 

needs. Evaluate and prioritize 

annually. 

E) Develop Mt. Lion population 

management methods that are 

consistent with established goals and 

objectives. 

F) Identify and describe suitable habitat 

areas and parameters for Mt. Lions 

in SD by Sept. 2003. 

G) Develop a comprehensive Public 

Education strategy for informing and 

educating the Staff, citizens and

44 SOUTH DAKOTA MOUNTAIN LION STATUS REPORT · Kintigh 

visitors about Mt. Lions and personal 

safety while in Mt. Lion country. 

H) Develop a public involvement plan 

for implementation during 2003 and 

2004 for inclusion in our 

management planning process. 

Over the last 10 years South Dakota has 

not significantly changed the way we 

manage lions. During this period of time 

they remained on the State’s Threatened 

Species List and very little was done to 

manage them other than offering them full 

protection of the law. Our awareness of 

lions did increase significantly during this 

time as we observed a steady apparent 

increase in their numbers. In recent years an 

Action Plan was developed to guide staff in 

dealing with problem lions. This Action 

Plan is currently under revision and will be 

included in the overall Management Plan. 

We also created a system for documenting 

and tracking lion activity. More significant 

changes are looming on the horizon as we 

remove the lion from the Threatened Species 

List and focus on concerted effort to manage 

our lions. 


Lions are currently distributed 

throughout the Black Hills, which contains 

the most suitable habitat in South Dakota. 

Some evidence of breeding populations also 

exists in the Custer National Forest in 

Harding County, the Badlands of eastern 

Pennington County and on the Pine Ridge 

Reservation of Shannon, Jackson and Bennett 

counties. 

Reports of lion activity have been 

received across all of South Dakota. 

Verification of reports outside of the Black 

Hills has proven to be very difficult, 

especially east of the Missouri River. Most 

occurrences outside of the Black Hills have 

been associated with river drainages that 


provide marginal habitat. 

The lion population in South Dakota 

appears to be still growing slowly at this 

time. Some uncertainty exists as to what the 

carrying capacity of the Black Hills for lions 

may be, though it is generally felt we are 

very close to that level now. Some evidence 

of dispersal from the Black Hills exists. To 

date we have only detected young males 

dispersing from the Black Hills. One young 

female was radio collared on the very edge 

of the Black Hills and some thought that she 

might disperse was expressed. She was 

poached before that determination was 

made. 

The cougar population in the Black Hills 

was estimated using program PUMA (Beier 

1993), incorporating parameters obtained 

from radio-collared cougars and habitat 

quality derived from a habitat-relation 

model. Annual home ranges were generated 

for 10 adult cougars monitored > 8 months, 

and spatial distribution of established males 

was analyzed using a home range overlap 

index. The area of the Black Hills was 

estimated at 8,400 km 2 , comprised of 

6,702.9 km 2 of high quality and 1,697.1 km 2 

of lower quality habitat (based on a habitatrelation 

model developed for the species). 

Mean annual home range size of established 

adult male cougars (n = 3) was 809.2 km 2 , 

and was significantly larger (P < 0.05) than 

that of adult females (n = 4), 182.3 km 2 . 

Based on sightings of family groups and 

radio-collared females, we documented up 

to 5 females occurring in established male 

ranges. Percent overlap for 3 established 

cougars averaged 33% (range = 18.0 - 

52.0%). Based on 5 population simulations, 

the total number of cougars in the Black 

Hills was estimated to be 127 to 149 

cougars; 46 to 49 adult females, 12 to 29 

adult males; 21 to 24 yearling females and 

males; and 45 to 48 female and male kittens.


South Dakota has not had any form of 

legalized mountain lion hunting since 1978. 

The future management of lions in South 

Dakota will include consideration of a 

hunting season as a management tool. 

Concerns about the impacts of hunting to the 

stability of the population will weigh heavily 

when those decisions are made. 



South Dakota does operate with an 

“Action Plan For Managing Mountain 

Lion/Human/Property Interactions.” An 

Action Plan was first developed in May of 

1995 and has been revised since then. This 

plan is included in the overall Mountain 

Lion Management Plan, which is currently 

under development. For our agency, 

addressing “problem” lions is the most 

difficult aspect of maintaining a population 

of lions. Public emotions are strong and 

varied which results in many 

comments/opinions being expressed directly 

at the “Action Plan.” 

South Dakota’s Action Plan categorizes 

Human/Lion interactions into five types: 

1. Sighting - a visual observation of a 

lion or a report of lion tracks or other 

sign on unpopulated lands or rural 

areas within the Black Hills. 

2. Encounter - an unexpected direct 

neutral meeting between a human and 

SOUTH DAKOTA MOUNTAIN LION STATUS REPORT · Kintigh 45 


a lion without incident (Mountain lion 

sightings in close proximity to homes, 

stables or livestock in rural areas and 

unpopulated lands outside of the 

Black Hills). A mountain lion is 

observed for the first time in close 

proximity or within residential 

developments and occupied 

recreational area. 

3. Incident - a conflict between a human 

and lion that may have serious results 

(e.g. a lion that must be forced to back 

down). Recurring observations of a 

lion in close proximity or within 

residential developments and 

occupied recreational areas. 

Livestock is killed in rural areas. 

4. Substantial public threat - a mountain 

lion that is observed within a city near 

areas where children are regularly 

congregated, killing wildlife/pets 

residential developments or occupied 

recreational areas or repeatedly killing 

livestock. 

5. Attack - when a human is bodily 

injured or killed by contact with a 

mountain lion. 

Each occurrence requires an 

understanding of all the circumstances and 

any history involved before an action is 

decided upon. In general, with every report 

of a lion a field investigation is highly 

encouraged by agency personnel (Table 1). 

Verification is key to any response. 

Table 1. Public Safety reports and resulting lion removals in South Dakota, 1998 – 2002. 

Year Number 

Reports 

Number 

Incidents 

Number 

Encounters 

Threatening 

Encounters 

Number of Public 

Safety Incidents 

Number Lions 

Removed 

1998 57 5 2 2 5 0 

1999 54 1 0 0 1 0 

2000 66 5 4 1 1 1 

2001 144 4 8 3 4 0 

2002 198 5 6 2 2 1


Table 2. Mountain lion depredations, verified depredations, and resulting lion removals in South 

Dakota, 1998 – 2002. 

Year Number Depredations 

Number Depredations 

Verified 


Number Lions 

Removed 

1998 1 1 0 

1999 0 0 0 

2000 2 1 0 

2001 3 2 1 

2002 4 2 0 

Note – one lion has been removed due to livestock depredation in 2003 already. 

Personnel are encouraged to take every 

opportunity to educate the public regarding 

all aspects of living with lions. Each lion 

reporting person receives an agency 

produced brochure on Mountain Lions. 

Public education is emphasized at this time 

and every opportunity is taken. 

Keeping all options available to 

responding staff is very desirable to our 

agency. However, we will not pay for any 

damages incurred due to wildlife of any 

species. 

Relocation of problem lions was once 

considered, but, due to the geographically 

limited area of the Black Hills and the 

existing lion population, it has been deemed 

an option that was unlikely to produce 

desirable results. Unusual circumstances 

may arise in which it may be attempted and 

the option has not been made totally 

unavailable. 

In rare cases, usually involving a single 

livestock producer, a permit has been issued 

for that individual to kill a lion that has been 

causing livestock depredation. Usually this 

only happens after agency efforts to remove 

the offending lion have failed. 

Our agency is equipped with a trio of 

trained lion hounds managed by an 

experienced houndsman. In most situations 

that necessitate a lion removal, the action is 

lead by our houndsman. Our state trappers 

are also equipped with leg snares, which are 

generally only set around livestock kills as 

the houndsman prepares to arrive on scene. 

On a few occasions, when a lion was a 

concern, but did not warrant removal we 

have chased the lion with hounds to haze the 

lion. On at least one occasion the lion was 

treed and a radio collar was fitted to increase 

our knowledge of its activity. 

In regards to livestock depredation, we 

currently investigate every report of this but 

take slightly different approaches to 

resolution depending upon the location. 

Livestock kills within the Black Hills 

typically require multiple kills before action 

to remove the offending lion is initiated. 

We are hesitant to remove lions from the 

limited quality habitat available in South 

Dakota (Table 2). Livestock depredation 

complaints on the plains of South Dakota, 

where limited habitat and a strong 

agricultural industry exists, are addressed 

much more decisively and quickly. 


The Department of Wildlife and 

Fisheries Sciences at South Dakota State 

University is currently completing a 5-year 

research project on cougars in the Black 

Hills. The main objectives of the research 

were to 1) develop and evaluate a cougar 

habitat-relation model to predict the current 

distribution 2) estimate the population size, 

and evaluate survey techniques to document

population trend. A digital habitat relation 

model was constructed for cougars that 

ranked land in the Black Hills National 

Forest according to its suitability to cougars. 

The model was based on the distribution of 

prey (white-tailed deer and mule deer), 

stalking topography (slopes), concealment 

habitat (riparian habitat), and anthropogenic 

characteristics (high-density residential 

areas, presence of highways). During the 

winters of 1998 – 2001, we captured, radiocollared, 

and obtained weekly locations of 

12 cougars in the Black Hills; locations of 

cougars were used to validate the habitatrelation 

model. The cougar population in 

the Black Hills was estimated using program 

PUMA, incorporating parameters obtained 

from radio-collared cougars and habitat 

quality derived from the habitat-relation 

model. The total number of cougars in the 

Black Hills was estimated to be 127 to 149 

cougars. 

A 3-month pilot study, testing the 

efficacy of detecting cougars using scent 

lures (skunk essence, Powder River cat call) 

and camera stations was conducted in 

cooperation with the University of North 

Dakota. The camera-scent-station survey 

was not effective at detecting cougar 

presence. Zero photos of cougars were 

recorded although other species (whitetailed 

deer, Odocoileus virginianus, mule 

deer, O. hemionus, raccoon, Procyon lotor, 

red squirrel, Tamiasciurus hudsonicus, 

turkey vulture, Cathartes aura, free-ranging 

cattle, feral dogs, and bobcat, Lynx rufus) 

were detected, and cougars were known to 

be in the area during the survey. A snowtracking 

helicopter survey (Vansickle and 

Lindzey 1991) using a probability sampling 

technique was attempted during the winter 

of 2001-2002. Although cougar tracks of a 

radio-collared female and her 2 kittens could 

clearly be identified, weather conditions 

(poor snow conditions) did not permit the 

survey to be completed. However, a 

SOUTH DAKOTA MOUNTAIN LION STATUS REPORT · Kintigh 47 


database of consecutive winter daily 

locations of 3 male and 3 female cougars 

was established to aid in analyses of any 

future helicopter surveys. 

During the Fall 2002, a second 5-year 

study was initiated. The objectives of the 

research are 1) to estimate survival and 

document causes of mortality of cougar 

kittens, 2) Determine longevity of 

established radio-collared cougars 3) 

Document dispersal distances, routes, and 

destinations of subadult cougars, and 4) 

conduct snow tracking helicopter population 

survey to document population trends. 

Currently, 12 cougars (6 females, 6 males) 

including 2 subadult males are being 

monitored weekly from fixed wing aircraft 

using aerial radio-telemetry techniques. 

PUBLICATIONS 

FECSKE, D.M., J.A. JENKS, AND F.G. 

LINDZEY. 2001. Characteristics of 

mountain lion mortalities in the Black 

Hills, South Dakota. Proceedings of the 

6th Mountain Lion Workshop, San 

Antonio, Texas: In Press. 

FECSKE, D.M., AND J.A. JENKS. 2001. The 

mountain lion returns to South Dakota. 

South Dakota Conservation Digest 

68(4):3-5. 

FECSKE, D.M., AND J.A. JENKS. 2001. 

Status report of mountain lions in South 

Dakota. Proceedings of the 6th 

Mountain Lion Workshop, San Antonio, 

TX. In Press. 

FECSKE, D.M., J.A. JENKS, AND F. G. 

LINDZEY. 2003. Mortality of an adult 

cougar due to a forest fire in the Black 

Hills. The Prairie Naturalist 00: 

Submitted. 

GIGLIOTTI, L.M., D.M. FECSKE, AND J.A. 

JENKS. 2002. Mountain lions in South 

Dakota: A public opinion survey. South 

Dakota Department of Game, Fish, and 

Parks, Pierre, SD. 182 pp. 

LONG, E.S., D.M. FECSKE, R.A. SWEITZER, 

J.A. JENKS, B.M. PIERCE, AND V.C.


BLEICH. 2003. Efficacy of photographic 

scent stations to detect mountain lions. 

Western North American Naturalist 00: 

In Press. 


BEIER, P. 1993. Puma: a population 

simulator for cougar conservation. 

Wildl. Soc. Bull. 21:356-357 


VAN SICKLE, W.D., AND F.G. LINDZEY. 

1991. Evaluation of a cougar population 

estimator based on probability sampling. 


743.

MOUNTAIN LION STATUS REPORT FOR TEXAS 

JOHN YOUNG, Texas Parks and Wildlife Department, 3000 IH 35 South Suite 100, Austin, TX 

78612, USA, email: john.young@tpwd.state.tx.us 

Texas does not currently have a 

statewide management plan for mountain 

lions and the species is classified as nongame. 

Texas Parks and Wildlife 

Department (TPWD) non-game codes 

authorize the agency to establish hunting 

seasons, to close seasons, set bag limits, 

establish management zones, in other words, 

to utilize all of the management tools 

available for game species. With the 

exception of a short list of non-game species 

of concern to TPWD, non-game species may 

be taken at any time of the year in any 

numbers, which is the case for mountain 

lions at the present time. TPWD’s objective 

for mountain lions is to maintain a viable 

population, while minimizing human 

conflicts. No changes in mountain lion 

status have occurred in the past decade. 


Based on confirmed sightings and 

mortality records mountain lions are most 

common in the Trans Pecos and the brush 

country of South Texas. Mortality records 

49 


over the last 20 years combined with photos 

confirm at least the occasional presence of 

lions in all other sections of the state; more 

information is needed to determine 

population levels. Based on sightings, and 

voluntarily reported mortalities dating back 

to 1983, mountain lion populations appear 

stable. Table 1 presents mountain lion 

mortality information by ecological region 

for the time frame 1998/99 to 2001/02 

Texas does not currently estimate mountain 

lion populations, opting to monitor the 

species using sightings and mortality 

reports. The lack of a satisfactory 

scientifically rigorous method to estimate 

mountain lions has been the primary reason 

TPWD has not attempted to do so. Texas 

has recently provided funding to a 

university-based scientist to estimate 

mountain lion population size, structure, and 

habitat factors utilizing new, highly credible 

molecular genetics. The study will be 

conducted over the next 2 years and will 

provide an estimate for Texas’ mountain 

lion population. 

Table 1. Mountain lion mortalities by ecological region, September 1998 through September 2002. 

Ecological Region 1998/99 1999/00 2000/01 20001/02 

Pineywoods 0 0 0 0 

Gulf Prairie & Marshes 0 3 0 0 

Post Oak Savannah 0 0 0 0 

Blackland Prairies 0 0 0 0 

Cross Timbers 0 0 0 1 

South Texas Plains 7 10 0 4 

Edwards Plateau 30 14 6 12 

Rolling Plains 0 0 0 0 

High Plains 0 0 0 0 

Trans-Pecos 92 60 64 48

50 TEXAS MOUNTAIN LION STATUS REPORT · Young 


Texas relies primarily on hunters, private 

landowners, and trappers to voluntarily 

report mountain lion kills. Texas also 

obtains an annual report from Texas 

Wildlife Damage Management Services 

(Table 1). There is an open season on 

mountain lions in Texas year-round. TPWD 

does not set harvest guidelines or bag limits 

for this species. Mountain lions may be 

taken by trap, shooting, hunting with dogs, 

aerial hunting, or M44. Records on the 

number of lions harvested by different 

methods are not collected. 

TPWD does not have a predator incident 

manual/policy/guideline for mountain lions 

although such has been developed for black 

bear. In the past 10 years there are only 3 

known public safety incidents in Texas 

related to mountain lion. Due to their rarity, 

TPWD does not formally record/collect 


information on public safety incidents 

involving mountain lion. Depredation 

complaints received at TPWD are referred to 

Texas Wildlife Damage Management 

Services (TWDMS). In 2001/02 a total of 

53 lions were killed by TWDMS personnel. 

Information on cougars removed by 

TWDMS prior to 2001/02 had been 

combined with other mortalities and has not 

been available separately. 

Individuals wishing to report a sighting 

or a problem with mountain lions are 

encouraged to contact TPWD. The 

department provides individuals 

experiencing depredation problems with the 

number for their local TWDMS office for 

action. Relocation of mountain lions is 

discouraged but may be conducted by 

private organizations if they acquire the 

appropriate permits.

UTAH MOUNTAIN LION STATUS REPORT 

CRAIG R. McLAUGHLIN, Utah Division of Wildlife Resources, 1594 W. North Temple, Salt 

Lake City, UT 84114, USA, email: craigmclaughlin@utah.gov 

Abstract: Mountain lions have been managed as a protected game species in Utah since 1967. In 1999 the Division 

of Wildlife Resources completed the Utah Cougar Management Plan, developed with the assistance of a publicbased 

Cougar Discussion Group that will guide management of cougars through 2009. Cougar harvests are 

managed under both harvest objective (quota) and limited entry strategies. The Division manages to sustain cougar 

densities on all management units except those that have approved predator management plans, where cougar 

harvests are increased to reduce cougar numbers and predation on big game. All cougar complaints are handled 

under the guidance of a Nuisance Cougar Complaints policy. Most cougar conflicts are handled through lethal 

control. Cougar habitat encompasses about 92,696 km 2 (35,790 mi 2 ). The statewide population was estimated at 

2,528-3,936 cougars in 1999 in conjunction with the Cougar Management Plan. Cougar harvests have ranged from 

492 to 373 annually since the 1997-1998 season. Both the hunting and pursuit seasons run from mid-December 

through June, although some units have extended or shortened seasons. Cougars have been implicated in 74-114 

separate depredation incidents per year since 1998, with livestock losses ranging from $53,700 to $97,700 per year. 

Harvest-based indicators of sustainable harvesting have not been met in recent years. Currently, management is 

operating on an individual-unit scale, where interpretation of harvest data is hampered by small sample sizes. In 

addition, the Division should develop a means to monitor both reproduction and survival. Harvest management 

should improve with understanding of cougar movements and dispersal, particularly between lightly hunted and 

heavily harvested cougar populations. 

51 


Key words: Cougar, livestock damage, harvest, management plan, mountain lion, Puma concolor 

INTRODUCTION 

Mountain lions (Puma concolor), or 

cougars, were persecuted as vermin in Utah 

from the time of European settlement (in 

1847) until 1966. In 1967 the Utah State 

Legislature changed the status of cougars to 

protected wildlife and since then they have 

been considered a game species with 

established hunting regulations. The Utah 

Division of Wildlife Resources (UDWR) 

developed the Utah Cougar Management 

Plan in 1999 (UDWR 1999b) with the 

assistance of a Cougar Discussion Group 

composed of representatives of various 

public interest groups. This plan will guide 

cougar management in Utah through 2009. 

Its goal is to maintain a healthy cougar 

population within existing occupied habitat 

while considering human safety, economic 

concerns and other wildlife species. 

Management objectives include: 1) 

maintaining current (1999) cougar 

distribution, with a reasonable proportion of 

older age animals and breeding females, 

balancing population numbers with other 

wildlife species; 2) minimizing the loss in 

quality and quantity of existing critical and 

high priority cougar habitat; 3) reducing the 

risk of loss of human life and reducing 

chances of injury by cougar; 4) maintaining 

a downward trend in the number of livestock 

killed by cougar; and 5) maintaining quality 

recreational opportunity for a minimum of 

800 persons per year through 2009. 

Utah’s cougar harvests are controlled on 

specific geographic areas, or management 

units (Figure 1), using two harvest 

strategies: harvest objective and limited 

entry. Under the harvest objective 

strategy, managers prescribe a quota, or 

number of cougars to be harvested on the 

unit. An unlimited number of licensed

52 UTAH MOUNTAIN LION STATUS REPORT · McLaughlin 

Figure 1. Wildlife Management Units used 

by Utah Division of Wildlife Resources to 

manage cougar harvests. Some of the units 

have been subdivided for additional control 

of harvests. 

hunters are allowed to hunt during a season 

that is variable in length, as the hunting 

season closes as soon as the quota is filled or 

when the season end date is reached. Under 

the limited entry strategy, harvests are 

managed by limiting the number of hunters 

on a unit. The number of hunters is 

determined based upon an expectation of 

hunting success and the desired harvest size. 

Individuals are usually selected for hunting 

on the unit through a random drawing 

process. 

In 1996 the Utah Wildlife Board 

approved a Predator Management Policy 

(UDWR 1996) that allows UDWR to 

increase cougar harvests on management 

units where big game populations are 

depressed, or where big game has recently 

been released to establish new populations. 

Most predator management plans directed at 

cougars have been designed to benefit mule 


deer (Odocoileus hemionus) and bighorn 

sheep (Ovis canadensis). Cougar harvests 

have been liberalized where big game 

populations are far below objective (

Figure 2. Cougar habitat in Utah. All 

colored areas represent occupied cougar 

habitat. 

Wasatch Mountains in northern and central 

Utah. 

The last statewide cougar population 

estimates were developed in conjunction 

with the Utah Cougar Management Plan in 

1999 (UDWR 1999b). These estimates used 

extrapolations of cougar densities from 

published studies in the southwestern United 

States to: 1) the total area within all 

management units that comprise cougar 

range, and 2) the total amount of occupied 

cougar habitat within Utah. The habitat 

quality within each management unit was 

classified as either high, medium or low 

based on vegetative characteristics, terrain 

ruggedness (following Riley 1998) and prey 

density. Cougar densities derived from 

research within Utah, California and New 

Mexico were associated with each habitat 

quality level (UDWR 1999b). High quality 

habitat was assigned a density range of 2.5- 

3.9 cougars/100 km 2 , medium quality 

UTAH MOUNTAIN LION STATUS REPORT · McLaughlin 53 


habitat was assigned a density of 1.7-2.5 

cougars/100 km 2 and a density of 0.26-0.52 

cougar/100 km 2 was assigned to low quality 

habitat. 

The first statewide population estimate 

of 2,528-3,936 cougars resulted from 

summing unit population estimates. The 

number of cougars on each unit was 

estimated by first multiplying the total area 

contained within the unit by the highest 

density of the range assigned to it, and then 

by the lowest density of the range assigned 

to it. 

For comparison, a second estimate of 

2,927 cougars statewide was generated 

based upon mean cougar densities and total 

occupied cougar habitat within the state. 

Each management unit’s cougar population 

was estimated by extrapolating the mean 

cougar density assigned to the unit (based on 

the respective range indicated above) to the 

amount of occupied cougar habitat within 

the unit, and unit estimates were summed to 

obtain the statewide figure. The two 

methods produced population estimates that 

show considerable agreement, but they 

should be only viewed as general 

approximations of the statewide cougar 

population. 

Utah’s cougar population is monitored 

through mandatory reporting of all hunterharvested 

cougars, cougars that are killed on 

highways or in accidents and those taken by 

animal damage control programs (Table 1). 

Location of kill, sex and age (through a 

premolar for age estimation) are recorded 

for every cougar killed, and provide the data 

used to assess management performance in 

relation to established target values that 

serve as indicators of population status. 

“Rules of thumb”, expressed as threshold 

values of 1) a minimum percentage of older 

aged animals in the harvest, 2) a maximum 

percentage of females in the harvest, and 3) 

minimum adult survival were set to ensure 

that cougar densities are maintained within


Table 1. Utah cougar harvests, 1989-1990 thru 2001-2002. 

Total Percent Percent 

Percent Treed 

Hunters Harvest Adult Adult Sub-adult Sub-adult Sport Average Quota Percent Female+ ADC Other Total Adult > per 

6 years Pursuit 

Year Afield Permits Objective Males Females Males Females Harvest Age Success Filled Females Sub-adult Harvest Mortality Mortality Survival old Day 

1989-90 478 527 123 44 23 27 217 41.2% 32.7% 43.3% 48 10 275 0.41 

1990-91 480 525 144 46 40 35 265 50.5% 30.6% 45.7% 38 22 325 0.49 

1991-92 485 525 128 51 32 30 241 45.9% 33.6% 46.9% 34 22 297 0.45 

1992-93 598 591 206 64 54 48 372 62.9% 30.1% 44.6% 53 42 467 0.49 

1993-94 575 659 165 87 51 49 352 53.4% 38.6% 53.1% 53 10 415 0.57 

1994-95 656 791 205 103 57 66 431 54.5% 39.2% 52.4% 54 24 509 

1995-96 787 872 160 105 109 78 452 3.5 51.8% 40.5% 64.6% 33 39 524 0.67 16.7% 0.48 

1996-97 1376 595 275 172 172 125 107 576 3.8 56.0% 88.3% 48.4% 70.1% 40 50 666 0.67 20.0% 0.33 

1997-98 1370 509 270 204 159 57 72 492 3.2 54.4% 79.6% 47.0% 58.5% 27 23 542 0.63 14.5% 0.36 

1998-99 1201 446 230 156 100 50 67 373 3.1 49.0% 64.0% 44.8% 58.2% 13 1 387 0.62 10.1% 0.29 

1999-00 817 343 304 194 106 64 71 435 2.9 60.0% 81.0% 40.7% 55.4% 25 9 469 0.57 9.7% 0.28 

2000-01 272 371 165 127 77 80 449 3.3 52.0% 35.4% 46.1% 63.3% 73 20 542 0.63 12.8% 0.37 

2001-02 258 339 159 108 55 71 393 2.9 45.5% 59.5% 12 7 412 0.61 9.0% 

Total 2181 1272 794 801 5048 

Average 802.1 531.8 298.2 167.8 97.8 61.1 61.6 388.3 3.2 52.6% 69.7% 39.8% 55.1% 38.7 21.5 448.5 62.7% 13.3% 0.41 

Performance Targets: 40.0% 65.0% 15.0% 0.38 

54 UTAH MOUNTAIN LION STATUS REPORT · McLaughlin


Table 2. Confirmed livestock losses due to cougar depredation in Utah, FY1992 to FY2002. 

Total Cougar 

Fiscal Year 

Number of Confirmed Losses: 

Incidents Ewes Lambs Bucks Calf Goat Other 

Confirmed 

Losses 

Value 

Losses 

Taken by 

WS 

1992 103 175 745 0 4 0 922 34 

1993 114 263 722 1 2 0 988 $94,644.00 53 

1994 115 258 646 5 6 0 915 $120,615.00 53 

1995 152 335 760 24 12 0 1130 $111,495.00 54 

1996 112 257 621 2 6 0 878 $79,277.00 33 

1997 110 375 531 20 11 0 937 $106,210.00 46 

1998 114 253 506 19 13 0 805 $97,703.00 27 

1999 69 244 406 18 4 0 730 $92,945.00 11 

2000 82 160 371 2 15 0 548 $60,750.00 22 

2001 74 136 361 12 3 1 587 $61,395.00 18 

2002 95 167 453 18 11 2 1 652 $53,748.42 74 

TOTAL 1140 2623 6122 121 87 3 1 8957 $825,034.00 351 

all management units, except where predator 

management plans are in place. Threshold 

values of the harvest criteria were obtained 

from the literature and from past evaluations 

of cougar population dynamics in Utah. This 

approach is likely conservative, but it is 

justified based upon our limited knowledge 

of the abundance of deer and alternate prey 

in Utah (UDWR 1999b). Ongoing research 

on 2 study sites, under the direction of Dr. 

Michael Wolfe (Utah State University), is 

supplying comparative data on the dynamics 

of cougars subjected to varying levels of 

hunting harvest. This information should 

help the Division refine management criteria 

in the near future. The Division also 

monitors trends in numbers of cougar 

incident reports, which have fluctuated in 

recent years (Table 2). Attempts to reduce 

the number of cougar management units that 

are subject to predator management plans 

have met with little success, mostly due to 

continued drought and deteriorating range 

conditions. 



Cougar hunting in Utah is regulated on a 

management-unit basis to address 

differences in cougar densities, hunter 

access and management objectives. 

Annually, the composition of each unit’s 

harvest is compared to performance targets 

that were selected to maintain cougar 

densities: 1) maintain an average of 15% or 

greater of the harvest in older age classes 

(>6 years of age); 2) maintain total adult 

survival at or above 65%; 3) restrict the 

female component to


The harvest objective strategy is often 

used on units where managers want to 

ensure a substantial harvest. This strategy 

can result in hunter crowding and less hunter 

selectivity toward males, as many hunters 

take the first cougar they encounter. 

Consequently, the harvest may be weighted 

toward young animals and females. 

Conversely, limited entry hunts allow 

managers to spread hunting effort over a 

longer time period and shift harvesting 

pressure toward adult males. This strategy 

is commonly used on management units that 

are readily accessible to hunters to minimize 

crowding and promote hunter selectivity for 

adult males. 

Since 2001, a few units have been 

harvested under a hybrid strategy, where 

both harvest objective and limited entry 

hunts are held. This approach attempts to 

produce a large harvest while encouraging 

some hunter selectivity. Under the hybrid 

strategy, a limited entry hunt is opened 

early, followed by a harvest objective hunt 

that is delayed until mid-winter. In the past, 

managers have used female sub quotas in 

conjunction with harvest objective strategies 

to protect females in the face of increased 

harvest pressure. This strategy has been 

discontinued because it biased the harvest 

sex composition toward females (through 

early closure when the sub quota was 

attained) and prevented meaningful 

evaluations of harvest sex composition 

under criterion 3 above. 

Each year, regional wildlife managers 

review the size and composition of harvests 

from individual units in relation to 

management rules of thumb and then make 

recommendations for the forthcoming 

season. Often, their evaluations result in 

changes in the number of permits allocated, 

the size of quotas and/or changes in harvest 

strategy. These regulation changes often 

result in year-to-year fluctuation in harvest 

strategy and hence harvest pressure. As a 


result, variances in harvest size and 

composition are difficult to interpret. Total 

harvest has varied from 492 to 373 since the 

1997-1998 season, with no definite trend 

(Table 1). 

Nearly all cougars harvested in Utah are 

taken with the aid of dogs. An individual 

hunter is restricted to holding either a 

limited entry permit or a harvest objective 

permit per season, and must wait 3 years to 

reapply once he/she acquires a permit. The 

bag limit is 1 cougar per season and kittens 

and females accompanied by young are 

protected from harvest. Currently the 

cougar-hunting season runs from December 

14, 2002 through June 1, 2003 on both 

limited entry and harvest objective units. 

However, some units are open year-round 

and some have earlier or later opening dates. 

Because harvest objective units close as 

soon as the objective (quota) is reached, 

hunters must call a toll-free number daily to 

ensure that the season in their hunt unit is 

still open. 

Pursuit (chase or no-kill) seasons 

provide additional recreational opportunities 

over most of the State. The pursuit season 

generally runs December 14, 2002 through 

June 1, 2003, but specific units have yearround 

pursuit and a few units are closed to 

pursuit hunting. In recent years, the Division 

has sold about 600-700 cougar pursuit 

permits annually (Table 3). 

The Division began managing cougar 

harvests through statewide limited entry 

hunting in 1990 and increased numbers of 

Table 3. Number of cougar pursuit permits 

sold in Utah, 1999-2002. 

Year Resident Non-Resident Total 

1999-2000 572 49 621 

2000-2001 595 59 654 

2001-2002 621 84 705 

Combined 1788 192 1980


Table 4. Comparison of harvest characteristics for Utah management units that have predator 

management plans (designed to reduce cougar numbers) and units that are managed to sustain 

cougar populations. 

Criteria (Threshold for 

sustaining population) 

Predator Management Plan in Place 

1999-2000 2000-2001 2001-2002 

% Females ( 6 years (>15) 9.7 9.8 10 

Adult Survival (>0.65) 0.60 0.61 0.52 

Cougar treed/day (0.38) 0.24 0.16 

permits through 1995-1996 (Table 1). In 

1996-1997, additional harvest pressure was 

added by switching some management units 

to the harvest objective (quota) system and a 

record high of 1,376 hunters was afield 

(Table 1). Since then, the number of hunters 

afield has declined nearly one-third. The 

hunting harvest has declined over the same 

period (Table 1). 

Units with predator management plans 

designed to reduce cougar densities produce 

harvests of similar composition to areas 

where the management objective is to 

sustain higher population densities (Table 

4). Throughout the State, the proportion of 

harvest comprised of females has usually 

been above the prescribed threshold for 

maintaining cougar densities, the percent of 

older aged cougars in the harvest has 

remained below the desired threshold level, 

adult survival is below the desired level, and 

the cougar treeing rate is below the value 

ascribed as an indicator of secure population 

abundance. Given the relative abundance of 

de facto refugia for cougars in Utah 

(National Parks, wilderness and inaccessible 

tracts) and the species’ propensity to 

disperse long distances, current harvest 

prescriptions may not prove effective for 

attaining either of the State’s management 


No Predator Management Plan 

1999-2000 2000-2001 2001-2002 

38 46 47 

7.6 12.3 9 

0.59 0.61 0.62 

0.30 0.24 

objectives (maintenance of population 

density, or substantial reduction in 

population density). 

Evaluation of Harvest Information 

The harvest-based criteria used in Utah’s 

cougar management system are based upon 

published research, and represent the 

expectation of harvest statistics that are 

associated with sustained population 

densities. However, managers have not 

been able to fully meet all threshold values 

since the Cougar Management Plan was 

adopted in 1999. There may be several 

explanations for this difficulty, including the 

geographic scale of management actions and 

differences in the vital rates of cougar 

populations within Utah. 

The proportion of mature (>6 years of 

age) cougars in the harvest is used as an 

index of the presence of mature cougars in 

the underlying population. If this proportion 

declines below 15%, the management plan 

assumes that the harvest rate is 

unsustainable. However, scarcity of olderaged 

cougars in harvests could also result 

from light (sustainable) harvesting of a 

productive cougar population by 

nonselective hunters, where relatively few 

cougars are taken and the harvest is


composed of mostly subadults and youngeraged 

adults. 

The proportion of adult females in the 

harvest is assumed to increase with 

increasing harvest pressure, and the 

threshold level chosen for sustainability in 

Utah (>40%) is based upon research from 

several western states. However, managers 

are evaluating small management units, 

some containing

understanding with UDWR. Their reports 

are compiled on a fiscal year basis (and 

therefore numbers/year differ from those 

reported in Table 1), and confirm livestock 

losses ranging from $53,700 to $97,700 per 

year since 1998 (Table 2). Cougars were 

implicated in 74-114 separate depredation 

incidents per year during this period, killing 

548-805 sheep, cattle and goats annually 

(Table 2). 


UDWR is funding research conducted 

through the Utah State University, under the 

direction of Dr. Michael Wolfe. This 

research has been ongoing on two study 

sites since 1995, and is directed at 

determining means of quantifying cougar 

populations and evaluating the effects of 

harvesting on them. Field research is 

currently underway by David Stoner, MS 

candidate. 

Recent Publications 

MAXFIELD, BRIAN D. 2002. Utah cougar 

harvest report 1998-1999. Annual 



performance report, Fed. Aid Project No. 

W-150-R-8. Publ. No. 02-07, Utah Div. 

Wildlife Res., Salt Lake City. 38 pp. 

MAXFIELD, BRIAN D. 2002. Utah cougar 

harvest report 1999-2000. Annual 

performance report, Fed. Aid Project No. 

W-150-R-8. Publ. No. 02-08, Utah Div. 

Wildlife Res., Salt Lake City. 41 pp. 


RILEY, S.J. 1998. Integration of 

environmental, biological, and human 

dimensions for management of mountain 

lions (Puma concolor) in Montaina. Ph. 

D. Diss., Cornell Univ. 158 pp. 

UDWR. 1996. Predator Management 

Policy. Utah Div. of Wildlife Res. Salt 

Lake City. UDWR. 1999a. Nuisance 

Cougar Complaints. Policy No. 

W5WLD-5. Utah Div. of Wildlife Res. 4 

pp. 

UDWR. 1999B. UTAH COUGAR MANAGEMENT 

PLAN. UTAH DIV. OF WILDLIFE RES. SALT 

LAKE CITY. 60 PP.

WASHINGTON COUGAR STATUS REPORT 

RICHARD A. BEAUSOLEIL, Bear / Cougar Specialist, Washington Department of Fish and 

Wildlife, 3515 Chelan Highway, Wenatchee, Washington, 98801, USA 

DONALD A. MARTORELLO, Bear, Cougar, and Special Species Section Manager, 

Department of Fish and Wildlife, 600 Capitol Way North, Olympia, Washington, 98501, 

USA 

ROCKY D. SPENCER, Dangerous Wildlife Specialist, Washington Department of Fish and 

Wildlife, 42404 North Bend Way SE, North Bend, Washington, 98045, USA 

INTRODUCTION 

Cougar (Puma concolor) occur 

throughout most of the forested regions of 

Washington State, encompassing 

approximately 88,497 km 2 or 51% of the 

State (Figure 1). Cougar became a protected 

big game species in 1966 and hunting 

seasons and harvest limits were established 

under the management authority of 

Washington Department of Fish and 

Wildlife (WFDW). In 1967, the 

Washington State Legislature passed a bill 

establishing a tag system in Washington. In 

1970, WDFW began mandatory reporting of 

cougar kills and in 1979 inspection and 

sealing of cougar pelts was required for data 

collection. In the mid-1980’s WDFW began 

collecting cougar teeth for age analysis. 

Figure 1. Distribution of cougars (gray) and 

cougar management units in Washington. 

60 


Currently, the statewide cougar management 

goal is to maintain healthy, self-sustaining 

cougar populations within each cougar 

management unit (CMU), except CMU 9, 

while minimizing the number of negative 

human-cougar interactions. 

HUNTING SEASONS AND HARVEST 

TRENDS 

Cougar seasons have changed 

significantly over the last several years 

(Figure 2). During the November 1996 

general election, Washington voters passed 

Initiative 655 (I-655) that banned the use of 

hounds for hunting cougar and bobcat, and 

the use of bait and hounds for hunting black 

bear. In an effort to mitigate the anticipated 

decrease in cougar harvest (i.e., post I-655), 

permit-only seasons were replaced with 

general seasons, cougar seasons were 

lengthened from approximately 6 weeks to 7 

and one-half months, and bag limit was 

increased from 1 to 2 cougar/year. 

Legislation was also passed that provided 

the authority to the Fish and Wildlife 

Commission to establish reduced costs for 

cougar and black bear transport tags, which 

they did from $24 to $5 in 1996 (cougar tags 

can also be purchased as part of a big game 

package). The outcome of these strategies is 

that the number of hunters purchasing a 

cougar tag in Washington has increased 

from 1,000 to 59,000. As a result, annual

350 

300 

250 

200 

150 

100 

50 

General Seasons 

Dogs Allowed 

Kill report required 

WASHINGTON COUGAR STATUS REPORT · Beausoleil et al. 61 

1979 - 1986 

1987 - 1995 1996 - 2002 

Figure 2. Cougar season structure and harvest in Washington, 1979-2002. 

cougar harvest during post I-655 years has 

increased slightly; however, the composition 

of the harvest has changed dramatically. 

The majority of cougar harvested pre-I 655 

was done so with the aid of dogs, thus 

mostly males and older animals were taken. 

Since 1996, the majority of cougars are 

harvested either as opportunistic encounters 

by deer/elk and cougar hunters, or by using 

tracking and calling techniques. These 

harvest methods are not as selective as using 

dogs. Therefore, since 1996, hunters have 

harvested more females and younger 

cougars (see oral presentation titled Cougar 

Harvest Characteristics With and Without 

the Use of Dogs in this proceedings). 

POPULATION STATUS AND TREND 

ANALYSIS 

The status of cougar populations is 

currently estimated through computer 

population simulation models, harvest 

characteristics, and, to a lesser degree, 

trends in human-cougar interactions. 

Based on population reconstruction 

models, harvest age data, and statewide 

cougar habitat estimates (using GAP 

analysis), the cougar population in 

Permit Seasons 

Dogs Allowed 

I 655 


General Seasons 

Dogs Banned 

Washington is likely between 2,400–4,000 

animals, and cougar population size is likely 

declining in a few areas of the state. 

Typically, the status of local or regional 

cougar populations are monitored via hunter 

effort and success, median age data, and 

percentage of females in the harvest; but 

only when viewed over several years with 

consistent harvest methods. Due to the 

changes in harvest methods during the last 

several years (predominantly hound hunters 

during pre I-655 years versus entirely spotstalk 

hunters during post I-655 years), no 

reliable trend data exist to accurately assess 

regional cougar populations or exploitation 

levels. As such, new population monitoring 

efforts are beginning in 2003, where cougar 

density and adult female survival will be 

evaluated and monitored in key areas of the 

State. 

HUMAN CONFLICT 

Human-cougar interactions are managed 

through public education, capture-removal, 

depredation permits, and public safety 

cougar removals. Since 1995, WDFW has 

recorded information on human-cougar 

interactions. Of particular concern is the

62 WASHINGTON COUGAR STATUS REPORT · Beausoleil et al. 

To address human safety 

To protect threatened and 

endangered species 

To prevent loss of domestic 

animals 

To increase game populations 

0 20 40 60 80 100 

Figure 3. During a general public opinion 

survey, the percent of Washington 

respondents that supported reducing 

predator numbers for specific purposes 

(Duda et al. 2002). 

increasing trend in human safety incidents, 

and pet and livestock depredations. When 

Washington citizens were asked about their 

attitudes regarding cougars, over 80% 

responded that reducing predator numbers 

for public safety is acceptable (Figure 3). 

Recognizing the widespread scope of the 

issue and its importance to cougars and 

people in the future, current cougar 

management goals include maintaining 

sustainable cougar populations and reducing 

human-cougar interactions. In some cases, 

reducing cougar populations to a lower, but 

sustainable level may help achieve both of 

these goals (Table 1). Given the recent 

Confirmed complaints 

1000 

900 

800 

700 

600 

500 

400 

300 

200 

100 


0 

247 

495 

563 

927 

694 

936 

498 

378 

1995 1996 1997 1998 1999 2000 2001 2002 

Year 

Figure 4. Total confirmed cougar complaints 

in Washington, 1995-2002 (includes human 

safety and pet/livestock incidents). 

history of high human-cougar interactions, 

WDFW developed a special cougar removal 

process to address cougar densities in areas 

with high levels of human-cougar 

interactions. Under rules adopted by the 

Fish and Wildlife Commission, public safety 

cougar removals occurred in 17 Game 

Management Units from Dec 15 – Mar. 15, 

in both the 2001-2002 and 2002-03 seasons; 

in those seasons 109 and 76 cougar were 

identified for removal and licensed hunters 

Table 1. Cougar population objectives for each cougar management unit in Washington, 2002. 

CMU Geographic Area Population Objective 

1 Coastal Maintain a stable cougar population 

2 Puget Sound Reduce * cougar population to enhance public safety and protection of property 

3 North Cascades Maintain a stable cougar population 

4 South Cascades Maintain a stable cougar population 

5 East Cascades North Reduce * cougar population to enhance public safety and protection of property 

6 East Cascades South Maintain a stable cougar population 

7 Northeastern Reduce * cougar population to enhance public safety and protection of property 

8 Blue Mountains Maintain a stable cougar population 

9 Columbia Basin Unsustainable; not considered suitable cougar habitat 

* Implement cougar population reductions over a 3-year period and monitor annually.

emoved 67 and 54 animals, respectively 

(61% and 71% success rate, respectively). 

Confirmed human-cougar incidents 

decreased by 47% during the 2001 calendar 

year from 936 in 2000 to 498 and an 

additional 24% in 2002 to 378 (Figure 4). 

MANAGEMENT CONCLUSIONS 

The statewide cougar population appears 

to be declining at this time due to increased 

female harvest and objectives to address 

public safety and protection of property. 

Given the distribution of cougars in 

Washington and the projected growth of 

human populations, interactions between 

humans and cougars will likely continue. 

WASHINGTON COUGAR STATUS REPORT · Beausoleil et al. 63 


As such, the long-term future of cougar in 

Washington ultimately rests in our ability to 

co-exist. Therefore, management efforts 

should continue to look for ways to 

minimize human-cougar interactions, 

particularly at the local population level. 


DUDA, M.D., P.E. DE MICHELE, M. JONES, 

W. TESTERMAN, C. ZURAWSKI, J. 

DEHOFF, A. LANIER, S.J. BISSELL, P. 

WANG, AND J.B. HERRICK. 2002. 

Washington residents’ opinions on and 

attitudes toward hunting and game 

species management. Harrisonburg, 

Virginia, USA.

WYOMING MOUNTAIN LION STATUS REPORT 

SCOTT A. BECKER, Trophy Game Section, Wyoming Game and Fish Department, 260 Buena 

Vista, Lander, WY 82520, USA, email: Scott.Becker@wgf.state.wy.us 

DANIEL D. BJORNLIE, Trophy Game Biologist, Wyoming Game and Fish Department, 260 

Buena Vista, Lander, WY 82520, USA, email: Dan.Bjornlie@wgf.state.wy.us 

DAVID S. MOODY, Trophy Game Section Coordinator, Wyoming Game and Fish Department, 

260 Buena Vista, Lander, WY 82520, USA, email: Dave.Moody@wgf.state.wy.us 

INTRODUCTION 

Management of mountain lions (Puma 

concolor) has changed markedly since the 

nineteenth century. In 1882, the Wyoming 

Territorial government enacted legislation 

placing a bounty on mountain lions and 

other predators. This allowed for lion 

hunting throughout the year and no bag 

limits were enforced. In 1973, the mountain 

lion was reclassified as a trophy game 

animal, which made the Wyoming Game 

and Fish Department (WGFD) fiscally liable 

for confirmed livestock losses. The 

following year, the first hunting season was 

established that included the entire state as a 

single hunt area, a bag limit of 1 lion per 

year was enacted, kittens and females with 

kittens at side were protected, and hunters 

were required to present skulls and pelts of 

harvested lions to the nearest WGFD 

District Office or local game warden. 

In 1997, the WGFD prepared a draft 

management plan for mountain lions, but the 

plan has yet to be finalized. However, six 

main objectives outlined in the draft 

management plan continue to guide lion 

management objectives for the state of 

Wyoming, they are: 1) maintain mountain 

lion populations within suitable habitat 

throughout Wyoming; 2) provide mountain 

lion-related recreational opportunities; 3) 

minimize female lion harvest in areas where 

population stability or increase is desirable; 

4) minimize mountain lion depredation and 

64 


lion/human interactions; 5) tailor 

management objectives to conditions present 

within each Mountain Lion Management 

Unit (MLMU) where possible; and 6) 

implement more specific, quantifiable 

objectives within each MLMU as 

information on the state’s lion population 

allows. Using these objectives as 

guidelines, the WGFD attempts to balance 

recreational demand and harvest with the 

biological needs of lion populations 

throughout the state. 


Mountain lions are distributed 

throughout nearly all habitats in Wyoming 

although densities are not uniform. Lion 

densities are thought to be highest in the 

Bighorn, Owl Creek, and Laramie mountain 

ranges (Wyoming Game and Fish 

Department 1997), while some of the lowest 

densities may be found in the grasslands of 

northeastern Wyoming. In the Bighorn 

Mountains, Logan and Irwin (1985) found 

that mixed conifer and curl leaf mountain 

mahogany habitats were used most in 

relation to availability, whereas sagebrush 

grass habitat types were generally avoided. 

In the Snowy Range Mountains of 

southeastern Wyoming, lions were found at 

lower elevations during the winter and 

concentrated their use near the timber/prairie 

interface (Chuck Anderson, personal 

communication).

Figure 1. Mountain lion management units 

and hunt areas in Wyoming, 2002. 

WYOMING MOUNTAIN LION STATUS REPORT · Becker et al. 65 



Data on mountain lions are gathered 

annually among 28 hunt areas that are 

grouped into 5 MLMUs (Figure 1), the 

boundaries of which encompass large areas 

with contiguous topographic features and 

are believed to encompass population 

centers. Each hunt area has a maximum 

annual mortality quota that varies from 2 – 

34, with 5 areas also having a maximum 

female mortality quota (Table 1). If either 

quota is filled, the hunting season in that 

hunt area automatically closes. Currently, 

hunting seasons open on September 1 and 

close on March 31 for all hunt areas except 

Table 1. Wyoming mountain lion management units, hunt areas, season dates, and quotas for 

harvest year 2002. 

Mountain Lion 

Management Unit 

Northeast 

Southeast 

Southwest 

North-Central 

West 

Hunt Area Season Dates 

Annual Mortality 

Quota 

Annual Female 

Mortality Quota 

1 Sept. 1-Mar. 31 7 

24 Sept. 1-Mar. 31 2 

5 Sept. 1-Mar. 31 12 

6 Sept. 1-Mar. 31 34 

7 Sept. 1-Mar. 31 6 

8 Sept. 1-Mar. 31 8 

9 Sept. 1-Mar. 31 3 

25 Sept. 1-Mar. 31 3 

27 Sept. 1-Aug. 31 20 

10 Sept. 1-Mar. 31 6 

11 Sept. 1-Mar. 31 2 

12 Sept. 1-Mar. 31 6 3 

13 Sept. 1-Mar. 31 3 

16 Sept. 1-Mar. 31 6 

15 Sept. 1-Aug. 31 25 

21 Sept. 1-Mar. 31 25 

22 Sept. 1-Aug. 31 15 

23 Sept. 1-Mar. 31 15 8 

2 Sept. 1-Mar. 31 12 6 

3 Sept. 1-Mar. 31 8 4 

4 Sept. 1-Mar. 31 4 

14 Sept. 1-Mar. 31 9 

17 Sept. 1-Mar. 31 5 

18 Sept. 1-Mar. 31 12 

19 Sept. 1-Mar. 31 20 

20 Sept. 1-Mar. 31 15 

26 Sept. 1-Mar. 31 12 7 

28 Sept. 1-Mar. 31 3

66 WYOMING MOUNTAIN LION STATUS REPORT · Becker et al. 

15, 22, and 27, in which year round seasons 

exist. Quotas begin at the start of each 

hunting season and include all legal and 

illegal hunting mortalities. 

Mountain lion data in Wyoming are 

limited to information obtained annually 

from harvest or other documented forms of 

mortality. Since 1974, hunters have been 

required to present the skull and pelt of 

harvested lions to a district game warden or 

biologist at the nearest WGFD regional 

office within 72 hours after the harvest. 

Information collected during these 

inspections include: harvest date, location, 

sex, lactation status, estimated age, number 

of days spent hunting, whether or not dogs 

were used, and number of lions observed 

while hunting. Skulls and pelts must be 

presented in an unfrozen condition so teeth 

can be removed. Evidence of sex must 

remain naturally attached to the pelt for 

accurate identification. 

Legal shooting hours are from one-half 

hour before sunrise to one-half hour after 

sunset. The individual bag limit for lions is 

1 lion per hunter per calendar year, except 

for 1 hunt area in central Wyoming, where 1 

additional lion may be taken each calendar 

year. Kittens (

Total Lion Harvest 

250 

200 

150 

100 

50 

0 

78 

1993 

1994 

95 110 

145 144 

206 

214 

201 

186 

172 

1995 

1996 

1997 

1998 

1999 

2000 

2001 

2002 

Figure 2. Total Wyoming mountain lion 

harvest, 1993-2002. 

information into mountain lion harvest 

analyses in order to better assess mountain 

lion population trends. This will eventually 

aid in adjusting population objectives and, 

thus quotas, to ensure sustainable lion 

populations statewide. 

There has been a steady increase in 

harvest since 1993, which has leveled off in 

recent years at around 200 (Figure 2). Since 

1993, the average percent of females in the 

harvest has been 43%, ranging from 32% in 

1993 to 51% in 2000 (Figure 3). The 

percent of adults in the female harvest has 

steadily declined in the past 10 years, falling 

from around 70% adult females in 1993 and 

1994 to around 40% adults in 2001 and 2002 

(Figure 4). This decline in the past two 

years is likely due in part to a change in the 

criteria used to classify adults and juveniles 

prior to the 2001 hunting season. Since 

1993, hunter effort has ranged from 3.3 to 

5.8 days per lion for an average of 3.9 days 

per lion. Ninety-two percent of all 

successful hunters in Wyoming harvested 

lions with the aid of dogs from 1993 – 2002. 

DEPEDATIONS AND HUMAN-LION 


Currently, Wyoming uses a statewide 

protocol for managing trophy game 


Percent 

100% 

75% 

50% 

25% 


0% 

1993 

1994 

1995 

1996 

1997 

1998 

1999 

2000 

2001 

2002 

Percent Females Percent Males 

Figure 3. Percent male and female mountain 

lion harvest in Wyoming, 1993-2002. 

depredations and interactions with humans. 

A depredating lion is defined as a lion that 

injures or kills livestock or domestic pets. 

In addition, 4 types of human/mountain lion 

interactions are defined by the WGFD, they 

are 1) recurring sighting – repeated sightings 

of a particular lion; 2) encounter – an 

unexpected meeting between a human and a 

lion without incident; 3) incident – an 

account of abnormal lion behavior that could 

have more serious results in the future (e.g., 

a lion attacking a pet, or a lion exhibiting 

aggressive behavior, without attack, toward 

Percent 

100% 

75% 

50% 

25% 

0% 

1993 

1994 

1995 

1996 

1997 

1998 

1999 

2000 

Adult Females Juvenile Females 

2001 

2002 

Figure 4. Percent adult and juvenile female 

mountain lion harvest in the total female 

harvest in Wyoming, 1993-2002.


humans); and 4) attack – human injury or 

death resulting from a lion attack. Each 

incident is handled on a case-by-case basis 

and is dealt with accordingly based on the 

location of the incident, the threat to human 

safety, the severity of the incident, and the 

number of incidents the animal has been 

involved in. Every effort is made to prevent 

unnecessary escalation of incidents through 

an ascending order of options and 

responsibilities: 

1) No Management Action Taken 

- Informational packets are provided 

to the reporting party that describe 

mountain lion natural history and 

behavior, damage prevention tips, 

and what to do in the event of an 

encounter. 

2) Deterrent Methods 

- Removal or securing of attractant 

- Removal of depredated carcass 

- Removal or protection of livestock 

3) Aversive Conditioning 

- Use of rubber bullets 

- Use of pepper spray 

- Use of noise making devices or 

flashing lights 

- Informational packets provided to 

the reporting party 

4) Trapping and Relocation 

- If the above efforts do not deter the 

lion from the area, if public safety 

is compromised, if it is a first 

offense, of if it has been a lengthy 

span of time between offenses 



5) Lethal Removal of the Animal by the 

WGFD 

- If the above methods do not deter 

the lion, if public safety is 

compromised, or if the offending 

lion has been involved in multiple 

incidents in a short span of time 


- Wyoming statute 23-3-115 allows 

property owners or their employees 

and lessees to kill mountain lions 

damaging private property, given 

that they immediately notify the 

nearest game warden of the 

incident 

- Lions that have been removed from 

the population will be used for 

educational purposes 



Education is a very important aspect of 

human and mountain lion interaction 

prevention. Therefore, the WGFD works 

closely with hunters, outfitters, 

recreationalists, livestock operators, and 

homeowners in an attempt to minimize 

conflicts with trophy game animals. Every 

spring, the WGFD hosts bear and lion 

workshops throughout the state to inform the 

public about bear and lion biology, front and 

back-country food storage techniques, and 

what to do in the event of an encounter with 

a bear or lion. In addition, numerous 

presentations are given throughout the year 

to civic, private, and school groups. Media 

outlets are also used to inform, and in rare 

incidents warn, the general public about bear 

and lion safety issues and any recent 

sightings. 

Even with all the educational efforts 

undertaken by the WGFD and preventive 

measures taken by the public, conflicts with 

mountain lions do occur. The number of 

mountain lion conflicts have ranged from a 

low of 13 reported incidents in 2002 to a 

high of 64 reported incidents in 1997. There 

have been a total of 40 mountain lion/human 

interactions in Wyoming since 1996 with no 

major injuries or deaths reported. 

Wyoming statute 23-1-901 provides 

monetary compensation for confirmed 

livestock damage caused by mountain lions. 

The number of damage claims for the last 10 

years range from 11 in 1995 to 28 in 1998,

and payments made to claimants range from 

a low of $22,627 paid in 1999 to a high of 

$44,071 paid in 1998 (Table 2). One 

hundred percent of the mountain lion 

damage claims paid in 2002 was for sheep 

depredations. From 1996 to 2002, 84% of 

reported lion depredations in Wyoming have 

involved sheep, 6% have involved horses, 

6% unknown livestock species, and 4% have 

involved cattle. An average of 4.9 nuisance 

lions were removed annually in the last 10 

years while an average of 1 lion was 

translocated annually from 1996 – 2002 (no 

translocation data available prior to 1996). 

PUBLIC ATTITUDES 

In 1995, the WGFD contracted with the 

Survey Research Center at the University of 

Wyoming to determine attitudes and 

knowledge of Wyoming residents on 

mountain lions and mountain lion 

management (Gasson and Moody 1995). 

Over 71% of the approximately 500 



respondents believed lions were a benefit to 

Wyoming. Attitudes toward mountain lion 

hunting were generally supportive, with 

49.6% agreeing or strongly agreeing that 

mountain lion hunting should continue and 

29.3% disagreeing or strongly disagreeing. 

The remaining respondents were either 

neutral or did not answer. However, most 

(57%) disagreed or strongly disagreed that 

hunting lions with dogs should continue as a 

legal method of take. Only 25.3% of 

respondents agreed or strongly agreed, while 

the remaining respondents were neutral or 

did not respond to the question. A large 

majority of respondents (80.7%) agreed or 

strongly agreed that mountain lion hunting 

seasons should be modified to avoid 

harvesting kittens or running females with 

kittens. A large majority of respondents 

(71%) were also opposed to the use of dogs 

to run and tree lions during non-harvest, 

chase seasons. 

Table 2. Wyoming ten-year mountain lion damage claim and translocation/removal history (all 

causes). 

Year # Claims $ Claimed $ Paid Translocations Removals 

1993 29 33,214.56 30,002.53 

0 

1994 26 30,498.51 24,646.00 

a 

5 

1995 11 40,634.67 34,594.67 

a 

4 

1996 14 28,540.96 24,947.95 0 6 

1997 20 28,935.16 28,761.50 1 10 

1998 28 56,171.39 44,070.79 2 5 

1999 21 32,307.63 22,627.43 2 6 

2000 20 42,352.69 30,773.59 0 5 

2001 15 38,322.79 25,592.46 1 6 

2002 13 35,870.99 32,075.05 0 2 

Mean 19.7 36,686.74 29,809.20 0.86 4.9 

a 

No data available. 

a



ANDERSON, C.R., JR., AND F.G. LINDZEY. 

2003. Estimating cougar predation rates 

from GPS location clusters. Journal of 

Wildlife Management 67(2): 307-316. 

ANDERSON, C.R., JR., F.G. LINDZEY, AND 

N.P. NIBBELINK. In review. Estimating 

cougar abundance using probability 

sampling: an evaluation of transect 

versus block design. Journal of Wildlife 

Management 00(0): 000-000. 


In press. Monitoring changes in cougar 

sex/age structure with changes in 

abundance as an index to population 

trend. 

ANDERSON, C.R., JR., F.G. LINDZEY, AND 

D.B. MCDONALD. In press. Genetic 

structure of cougar populations across 

the Wyoming Basin: metapopulation or 

megapopulation. 


2000. A guide to estimating mountain 

lion age classes. Wyoming Cooperative 

Fish and Wildlife Research Unit, 

Laramie, Wyoming. 

GASSON, W., AND D. MOODY. 1995. 

Attitudes of Wyoming residents on 

mountain lion management. Planning 


rep. #40, Wyoming Game and Fish 

Department, Cheyenne. 7 pp. 

WYOMING GAME AND FISH DEPARTMENT. 

1997. Mountain Lion Management 

Plan. Wyoming Game and Fish 

Department. 30 pp. 


1999. Protocol for managing aggressive 

wildlife/human interactions. Wyoming 

Game and Fish Department. 17 pp. 


2003. Annual mountain lion mortality 

summary: harvest year 2002. Trophy 

Game Section, Lander, Wyoming. 22 

pp. 


GASSON W. AND D. MOODY. 1995. 

Attitudes of Wyoming residents on 

mountain lion management. Planning 

rep. #40, Wyoming Game and Fish 

Department, Cheyenne, 7 pp. 

LOGAN, K.A. AND L.L. IRWIN. 1985. 

Mountain lion habitats in the Bighorn 

Mountains, Wyoming. Wildlife Society 

Bulletin 13: 257-262. 


1997. Mountain Lion Management 

Plan. Wyoming Game and Fish 

Department. 30 pp.

CRYPTIC COUGARS - PERSPECTIVES ON THE PUMA IN THE EASTERN, 

MIDWESTERN, AND GREAT PLAINS REGIONS OF NORTH AMERICA 

JAY W. TISCHENDORF DVM, Director, American Ecological Research Institute (AERIE), 

Post Office Box 1826, Great Falls, MT 59403, USA, email: TischendorfJ@Hotmail.com 

Abstract: The subject of cougars in eastern North America continues to intrigue and perplex wildlife biologists, 

managers, and nature enthusiasts. Almost uniformly considered extirpated throughout states and provinces in 

eastern and midwestern North America over a century ago, growing numbers of reports, some accompanied by 

incontrovertible evidence such as full specimens, blood, scat, track, or film documentation, suggest that Puma 

concolor is re-establishing, or has re-established, itself in some areas of this vast region. Similar evidence exists for 

the Great Plains. This paper, while probably raising more questions than it answers, examines the best and most 

current evidence for the occurrence of cougars in the East, Midwest, and Great Plains; discusses the official status of 

the species; and provides a perspective on the scientific, social, and political opportunities and challenges posed by 

this fascinating and compelling situation. 

71 


Key words: cougar, recovery, East, Midwest, Great Plains, prairie, North America, Puma concolor 

INTRODUCTION AND OBJECTIVES 

The possible existence of the puma 

(Puma concolor) in eastern and midwestern 

North America today, approximately 100 

years since its supposed extirpation from the 

region, is among the most provocative and 

exciting mysteries in the modern realms of 

natural history, ecology, wildlife 

management, and conservation biology. 

Importantly, the story of the cougar in the 

East, the ghost of North America, as it was 

dubbed by Bruce Wright, an early champion 

for its recovery, has far-reaching, global 

implications for carnivore conservation, 

continental ecological equilibrium, and 

perhaps most of all, our own fulfillment as 

stewards of the planet (Wright 1959). To 

understand this yet unfolding story, several 

fundamental concepts need review: 

1. Throughout North America from the 

Great Plains eastward, with the 

exception of Florida, the puma was 

generally considered extirpated by the 

early 1900s (Young and Goldman 

1946). 

2. Since that time, in virtually every state 

and every province across this vast 

region, scores of people, including 

professional scientists, biologists, 

naturalists, and foresters, have been 

reporting observations of cougars or 

their sign (Wright 1972, Tischendorf 

and Henderson 1994). 

3. While many of these reports are 

unverifiable or erroneous, a surprising 

number have been confirmed, often 

with the details published in peerreviewed 

literature. This history of 

confirmed reports since the time of 

supposed extirpation suggests, at a 

minimum, the periodic presence of freeranging 

cougars in the region. 

4. Several plausible explanations exist for 

these cryptic cats: 1) continued 

existence of native pumas; 2) 

immigration of western cats; 3) 

presence of feral escaped or released 

captives (FERCs); or 4) combinations 

of any or all of these (Nowak 1976, 

Downing 1984).

72 CRYPTIC COUGARS · Tischendorf 

From ecological, social, and political 

standpoints there are three main questions 

that this paper seeks to answer. One, are 

there cougars in the aforementioned region 

today? Two, if pumas are present, do they 

represent a breeding population(s)? Finally, 

what is the future of Puma concolor east of 

the Rocky Mountains? What truly does the 

public want when it comes to large 

carnivore recovery or restoration in the 

East? Possibilities here include active 

recovery, passive recovery (i.e., the animals 

establish viable populations on their own 

without active, direct human intervention), 

or overt efforts to preclude recovery. 

DISCUSSION 

Puma Presence, Populations, and the Big 

Picture Perspective 

To effectively understand the cryptic 

cougar situation, it is critical to maintain a 

big picture perspective (Tischendorf 1992c, 

1996a, b). Among the many who have 

commented on the subject over the years, 

and especially among those skeptical of 

cougar presence or recovery east of the 

Rockies, this perspective, “from Nova 

Scotia to Nebraska” (Tischendorf 1996a:43), 

has often been lacking (Tischendorf 1992c; 

1996a, b). Such a perspective was, 

however, utilized by Bruce Wright and, 

more recently, by United States Fish and 

Wildlife Service (USFWS) researcher 

Robert Downing. Downing authored the 

eastern cougar recovery plan and speculated 

on the presence of an extremely low density, 

widely dispersed puma population in the 

eastern United States (USA) (Downing 

1981, 1984; United States Fish and Wildlife 

Service 1982). 

Downing’s views, coupled with updated 

range information synthesized by Allen 

Anderson and intensive independent review 

of 100 years’ worth of reports and 

documentation, led one author to 

subsequently suggest that there were 

actually upwards of four loosely interrelated 


puma populations in the East and Midwest 

(Downing 1981, 1984; Anderson 1983; 

Tischendorf 1993c). Each of these lowdensity 

puma populations was believed to 

consist of widely dispersed, widely roaming, 

and perhaps transient animals (Tischendorf 

1993c). These populations were believed to 

have their epicenters in the Canadian 

Maritimes-New England region, the Great 

Lakes-northern Midwest region, the 

Missouri-Arkansas-Oklahoma area, and the 

Southeast. 

This theory of course remains unproven, 

although it was revisited at a previous 

Mountain Lion Workshop by several of this 

author’s colleagues similarly associated with 

the West Virginia-based Eastern Cougar 

Foundation (ECF) (Bolgiano et al. 2000). 

Members of the ECF, formed in 1999, are 

utilizing automatic cameras in an attempt to 

document consistent cougar presence or 

family groups that could support the above 

hypothesis. The ECF (website at 

www.easterncougar.org) is notable in that it 

has been able to positively partner with 

several governmental agencies and share in 

the efforts to recover pumas in the East. 

Such critical cooperation is also 

demonstrated with the Eastern Cougar 

Network (ECN). This group’s website, 

www.easterncougarnet.org, is a nonadvocacy 

amalgamation of peer-reviewed 

contributions on the subject of cougars from 

essentially every state and provincial 

resource agency from the Great Plains 

eastward. The site thus serves effectively as 

a real-time source of scientifically based 

status information on the cat, and perhaps 

one day other predators including gray 

wolves (Canis lupus), black bears (Ursus 

americana), lynx (Lynx canadensis), and 

wolverines (Gulo gulo) east of the Rockies. 

Seemingly integral to the subject of 

cougars east of the Rockies is the question 

of whether the species persisted in its native 

state beyond the days of its supposed

extirpation. In the big picture, however, if 

free-ranging pumas are present and 

behaving in wild puma ways, then their 

origin, whether from native eastern or 

western stock or sanctioned or unsanctioned 

releases, should not alter their proper 

management and may be irrelevant. While 

the cats in many confirmed puma reports are 

written off as FERCs and denied 

consideration as legitimate ecological 

entities, the North American continent teems 

with a host of wildlife populations having 

their origins in captivity. These span the 

spectrum from critically endangered species 

to non-native exotics raised like barnyard 

fowl and annually introduced solely for 

sporting opportunities. Yet these former 

captives continue to benefit from official 

recognition, management, and protection. 

Should mountain lions that happen to show 

up in areas where their presence is 

considered improbable be any different? 

Having said this, the historically 

consistent pattern of sightings and periodic 

confirmations, while circumstantial, 

suggests native pumas did persist in many 

areas of their former midwestern and eastern 

range at least into the 1940s and 1950s. 

After World War II, however, ownership of 

cougars and other wild, exotic, or novelty 

animals became part of mainstream 

Americana and some captive cougars likely 

ended up as FERCs. Unfortunately this 

phenomenon continues today and is not 

necessarily limited to the eastern USA. As a 

result, the ultimate origin of almost any freeranging 

puma today, even with genetic 

testing, may truly be indeterminate. 

Summary of Occurrence Records 

In keeping with a big picture perspective 

of the cryptic cougar subject, it is useful to 

review a sampling of bonafide puma reports. 

Examples of confirmed or highly credible 

reports, mostly peer-reviewed, follow. 

“Confirmed kill” indicates that a puma was 

killed and the incident documented both 


CRYPTIC COUGARS · Tischendorf 73 

photographically and by written or verbal 

elaboration of substantial details, or without 

photos but with written or verbal elaboration 

of substantial details by a professional 

scientist or wildlife manager associated with 

or employed by a governmental natural 

resource agency or academic institution. 

“Reported kill” involves highly credible 

documentation by a natural resource 

professional of a mountain lion being killed, 

but reflects a lack of substantial details. 

“Confirmed tracks” indicates that a track or 

tracks consistent with those of a puma were 

located and documented via measurements 

and/or photographs subsequently published 

in mainstream scientific or popular literature 

and thus widely available for independent 

scrutiny and authentication. 

Reported kill - Williston, North Dakota, 

1902 (Bailey 1926) 

Reported kill - Bears Paw Mountains, 

Montana, 1910 (White 1967) 

Reported kill - Fontana Village area, 

Tennessee, 1920 (Linzey and Linzey 

1971) 

Confirmed kill - Mundleville, New 

Brunswick, 1932 (Wright 1972) 

Confirmed kill - Little Saint John Lake, 

Maine-Quebec border, 1938 (Wright 

1972) 

Confirmed kill - Madison, Saskatchewan, 

1939 (Clarke 1942) 

Confirmed kill - Pasquia Hills, 

Saskatchewan, 1948 (White 1963) 

Confirmed kill - Asheville, Alabama, 1948 

(Anonymous 1948) 

Confirmed kill - Mena, Arkansas, 1948 

(Lewis 1969, Nowak 1976) 

Confirmed kill - Sims, Arkansas, 1949 

(Sealander 1951) 

Confirmed kill - Black Hills, South Dakota, 

1958 (Mann 1959) 

Reported kills (2) - Newcastle, Wyoming, 

ca 1950s-1960s (Roop 1971) 

Reported kills (2) - Van Tassell, Wyoming, 

ca 1959-1960 (Roop 1971)


Confirmed kill - Keithville, Louisiana, 

1965 (Goertz and Abegg 1966) 

Confirmed kill - Edinboro, Pennsylvania, 

1967 (Doutt 1969) 

Confirmed carcass - Checotah, Oklahoma, 

1968 (Lewis 1969) 

Confirmed kill - Hamburg, Arkansas, 1969 

(Noble 1971) 

Reported kill - Ekalaka, Montana, ca 1970 

(Nowak 1976) 

Confirmed kill - Pikeville, Tennessee, 1971 

(Nowak 1976) 

Confirmed kill - Stead, Manitoba, 1973 

(Nero and Wrigley 1977) 

Confirmed kill - Cutknife, Saskatchewan, 

1975 (White 1976) 

Cougar reportedly trapped - Baca County, 

Colorado, 1976 (Boddicker 1980) 

Confirmed hematological evidence - 

Menominee County, Michigan, 1984 

(Bill Adrian, Colorado Division of 

Wildlife, personal communication) 

Puma trapped, radio-collared, translocated 

to Black Hills - central South Dakota, 

1990 or 1992 (Ted Benzon and Ron 

Sieg, South Dakota Department of 

Game, Fish and Parks, personal 

communication; Tischendorf and 

Henderson 1994) (Note: This cat was 

killed in the Black Hills in 1996 [Ron 

Sieg, South Dakota Department of 

Game, Fish and Parks, personal 

communication]) 

Confirmed kill - Golden Valley County, 

North Dakota, 1991 (Tischendorf and 

Henderson 1994) 

Confirmed kill - Pine Ridge area, Nebraska, 

1991 (Tischendorf 1992a, Tischendorf 

and Henderson 1994) 

Cougar trapped and translocated to 

Colorado - Worthington, Minnesota, 

1991 (Tischendorf 1992a, b) 

Confirmed kill - Lowery, South Dakota, 

1992 (Tischendorf and Henderson 

1994) 


Confirmed kill - Lake Abitibi, Quebec, 

1992 (Tischendorf 1993a) 

Confirmed tracks - McKiel Lake, New 

Brunswick, 1992 (Tischendorf 1993b, 

Cumberland and Dempsey 1994) 

Confirmed kill - Texas County, Missouri, 

1994 (Hardin 1996, Bolgiano et al. 

2000) 

Confirmed kill - Mitchell, Nebraska, 1996 

(Frank Andelt, Nebraska Game and 

Parks Commission, personal 

communication) 

Confirmed kill - Floyd County, Kentucky, 

1997 (Bolgiano 2001) 

Confirmed kill - Randolph County, Illinois, 

2000 (Clark et al 2002) 

Confirmed kill - Duluth, Minnesota, 2001 

(Anonymous 2002) 

Confirmed kill - Harlan, Iowa, 2001 

(Anonymous 2002, Clark et al 2002) 

Confirmed kill - Callaway County, 

Missouri, 2003 (Graham 2003) 

Almost 30 years ago Nowak (1976:143- 

144) commented, “The sum of evidence 

suggests that native cougar populations have 

maintained themselves in southeastern 

Canada, within the former range of F. c. 

cougar (sic: should be couguar), and in the 

Ozark Plateau and adjoining forests of 

Arkansas, southern Missouri, eastern 

Oklahoma, and northern Louisiana.” 

Indeed, even if ecologically significant 

populations did not persist, the above list 

suggests it is doubtful that these furtive 

felids were ever totally extirpated from the 

vast, and in many cases relatively 

inaccessible, environs of this area. 

Relatively pristine areas within New 

Brunswick, Quebec, Ontario, and Manitoba, 

for instance, could possibly have sustained 

individual pumas or even vestigial, remnant 

populations of these cats through the “Dark 

Age” of wildlife and habitat management 

late in the late 19 th and early 20 th centuries. 

In the USA, a number of areas could 

also have served as similar refugia. As late

as the mid-1940s, for instance, noted 

mammalogist Victor Cahalane 

acknowledged cougar presence in 

Shenandoah National Park and adjacent 

Blue Ridge country of Virginia (Cahalane 

1948). Other plausible refuges include 

northern Maine, the Adirondacks, the 

Quabbin Reservoir area in Massachusetts, 

northern Pennsylvania, the impenetrable 

southeastern and southern coastal swamps, 

the Great Smoky Mountains area, the 

Tennessee-Virginia-Kentucky-West 

Virginia border country, the dense 

northwoods of Michigan and Minnesota, the 

Ozark and Ouachita Mountain complex of 

Missouri, Arkansas, and Oklahoma, and the 

Black Hills of South Dakota and Wyoming. 

White-tailed deer (Odocoileus 

virginianus) and mule deer (Odocoileus 

hemionus) irruptions in many of these same 

areas were identified circa 1940, confirming 

that within only a few generations of the 

puma’s supposed demise these sites had an 

adequate prey base to sustain or attract the 

species (Leopold et al. 1947). While an 

alternative argument is of course that the 

irruptions resulted from lack of predators, 

irruptions were also noted in the Rocky 

Mountains, where historically we know 

mountain lion populations may have been 

depleted but were never decimated. 

Indeed, in seminal biological 

publications as late as the mid-1970s and 

1980s several noted mammalogists and 

wildlife scientists postulated the continued 

presence of puma populations in these very 

same areas (Cahalane 1964, Burt and 

Grossenheider 1976, Nowak 1976, Deems 

and Pursley 1978, Russell 1978, Hall 1981, 

Anderson 1983). 

Nonetheless, such reports seemingly 

generated merely passing interest from 

mainstream science and remained largely off 

the radar screen of wildlife biologists. 

Today, with the growing groundswell in 

conservation biology and corridor ecology, 



scientists are more inclined to recognize the 

importance and implications of the presence 

of small, insular predator populations or 

even remnant, wandering, or dispersing 

individuals. Indeed, such enclaves or 

individuals may provide the necessary seed 

for recolonization or recovery. 

Natural resource agencies responsible 

for some of these above-mentioned areas 

today, such as the Black Hills, have in fact 

confirmed extant puma populations 

(Tischendorf and Henderson 1994). 

Additionally, for quite some time other 

states such as North Dakota, Minnesota, 

Wisconsin, Missouri, and Arkansas have 

acknowledged at least limited and sporadic 

puma presence (Sealander and Gipson 1973, 

Gerson 1988, Tischendorf and Henderson 

1994, Clark et al. 2002, Graham 2003, 

Heisel 2003). Michigan Department of 

Natural Resources officials acknowledge the 

presence of pumas in the Great Lakes State 

as well, and ongoing work there at least 

suggests the possibility of a resident, 

breeding population (Johnson 2002, 

Zuidema 2002, Mike Zuidema, Michigan 

Department of Forestry, retired, personal 

communication). Given the habitat, cover, 

prey base, and presence of a thriving 

carnivore guild that includes populations of 

wolves, bears, coyotes (Canis latrans), 

bobcats (Felis rufus), fishers (Martes 

pennanti), and probably an occasional lynx, 

it would perhaps be more surprising if 

Michigan did not have a resident puma 

population. 

The situation in the Prairie Provinces of 

Canada, with their seemingly less 

sensational and less controversial approach 

to the cat, is similar, if not even more 

definitive. In Saskatchewan, the late Tom 

White documented the presence of a small 

population of pumas filtering among the 

coulees and more rugged reaches of this 

sprawling province (White 1982). The 

Yukon and Northwest Territories, and


Alaska as well, have a consistent history of 

credible puma reports, suggesting occasional 

dispersal, while Manitoba Conservation 

continues to recognize a stable and perhaps 

growing puma population in that province 

(Cahalane 1964; Weddle 1965; Kuyt 1971; 

White 1982; Wrigley and Nero 1982; Robert 

W. Nero, Manitoba Museum of Man and 

Nature, retired, personal communication). 

Cougar Comeback 

Some researchers believe that pumas, as 

wolves did in the northern Rocky Mountains 

in the 1980s, are in fact re-colonizing many 

areas in the Great Plains and central 

mountains eastward (Tischendorf and 

Henderson 1994). As is true for the Dakotas 

and Minnesota, most of the prairie states, 

including Nebraska, Kansas, Oklahoma, and 

Iowa acknowledge, if not resident then 

transient occurrences of pumas (Tischendorf 

and Henderson 1994; Johnson 1998, 2000). 

The same is true for the eastern portions of 

Montana, Wyoming, Colorado, and Texas, 

where, in some cases, sporadic puma 

presence has been noted for years but where 

documented occurrences of these “prairie 

panthers” are clearly on the increase 

(Boddicker 1980; Berg et al. 1983; Johnson 

1998, 2000; Riley 1991; Roop 1971; Russ 

1997). 

Deer-rich riparian zones along river 

systems such as the Yellowstone, Missouri, 

North and South Platte, Arkansas, Canadian, 

Red, and Colorado River in Texas, can 

undoubtedly serve as effective corridors for 

puma immigration all the way to the 

southeastern Texas coast, Mississippi River, 

and beyond. Additionally, the 

documentation of puma deaths along 

railroad tracks in Nebraska and Illinois 

suggests the possibility that railroad rightof-ways 

and associated brush belts may also 

be effective pathways for pumas (Frank 

Andelt, Nebraska Game and Parks 

Commission, personal communication; 

Clark et al. 2002). 


The same pattern of puma recolonization 

discussed above could be 

occurring from the mid-continent’s northern 

reaches south and eastward. For instance, 

Manitoba’s puma population may be linked 

with Ontario to the east and northern 

Minnesota, Wisconsin, and Michigan to the 

south. Conversely, if low numbers of pumas 

have in fact inhabited some of these areas all 

along, their acknowledged presence today 

may be a function of both immigration and 

numerical local growth. 

In the Northeast, a similar phenomenon 

of range reestablishment may be taking 

place. This sentiment was first voiced by 

Canadian biologist Bruce Wright, famed 

World War II frogman-commando, Leopold 

student, early champion for the eastern 

cougar, and a strong advocate for eastern 

carnivore recovery (Wright 1959, 1972; 

Tischendorf 1996a; Allardyce 2001). It was 

Wright’s belief that throughout European 

man’s settlement of the region panthers 

persisted in the central highlands of New 

Brunswick and by the mid-1900s were, like 

the spokes of a wheel, re-populating and 

reclaiming their former range in the East. 

This belief, while perennially difficult to 

reconcile with the lack of confirmed puma 

populations in New Brunswick, or anywhere 

else in the East outside of Florida, is 

exemplified by growing numbers of not 

simply puma reports, but of highly credible 

or even verified puma reports (Gerson 1988, 

Cumberland and Dempsey 1994, Snow 

1994, Stocek 1995, Bolgiano et al. 2000). 

These include specimens, scats, tracks, and 

videotapes depicting these cats across a wide 

geographical zone extending essentially 

from Ontario to Newfoundland and 

southward to Georgia. 

In the northeast USA, Maine and New 

York are perhaps the most promising in 

terms of numbers of credible puma reports. 

One rather compelling report from Maine 

involved a shaken hunter who, at extremely

close range, came upon the gripping scene 

of what he described as a puma mortally 

ravaging a bobcat (Tischendorf 1994a). A 

sampling of other data from Maine on file at 

the American Ecological Research Institute 

(AERIE) includes a hair sample 

confirmation from 1995, a track photograph 

from the mid-1990s, and a credible 2000 

report of what was thought to be a female 

puma and kitten. This author has also seen 

puma track photos taken by biologist George 

Matula of the Maine Department of Inland 

Fisheries and Wildlife at a deer yard during 

routine winter surveys circa 1984 or 1985. 

Credible New York puma reports on file 

with AERIE include the killing of a puma 

kitten by a bobcat hunter in the early 1990s 

and three believable sightings of individual 

pumas by professional natural resource 

workers. All of these events stem from the 

vast Adirondack Park area and occurred 

during the 1990s. 

Other areas of the Northeast are not 

without their own intriguing data. In 1994, 

for instance, a group of 3 pumas was 

observed and tracked near the community of 

Craftsbury, Vermont (Theodore Reed, 

Friends of the Eastern Panther, personal 

communication). Presumably an adult 

female with 2 kittens, a scat deposited by the 

group was collected; subsequent analysis 

confirmed presence of puma hairs (Bonnie 

Yates, USFWS Wildlife Forensic 

Laboratory, personal communication). A 

hair sample from the remote and 

untrammeled Gaspe’ Peninsula in northern 

Quebec was also recently confirmed as that 

of a puma by Marc Gauthier and his 

Canadian research team (Mark Dowling, 

Eastern Cougar Network, personal 

communication). 

Evidence of Breeding and Validity of 

Sighting Reports 

The questions of confirmed puma 

breeding and actual puma numbers are 

problematic. In the absence of systematic, 



scientifically objective, replicable, and 

typically expensive multi-year studies, such 

as those involving mark and recapture 

techniques or radio telemetry, it is difficult 

to extrapolate population-level 

characteristics of any animal. And, as 

several speakers at this conference have 

noted, even with robust, million dollar 

studies, it is difficult to quantify puma 

populations. Even more difficult and more 

expensive is monitoring a puma population 

over substantial periods of time. What does 

this bode for eastern and midwestern 

resource agencies trying to decode the issue 

of cryptic cats that many seem to report but 

few can verify? 

Complicating the issue is the fact that 

agencies and their human constituency east 

of the Rocky Mountains have limited 

exposure to large carnivores and their 

management. In this geographic area there 

truly is a different mindset and comfort level 

towards research and management involving 

these animals, especially those capable of 

attacking and killing people. In the Black 

Hills of South Dakota, for instance, radiotracking 

of a young male puma in the early 

1990s was discontinued after only a short 

time due to concerns over liability if the 

animal were implicated in damage to a 

human or to human property (Tischendorf 

and Henderson 1994). 

Again due to concerns over liability, 

Missouri officials are reluctant to approve 

any studies involving handling or marking 

of black bears, which are apparently 

repopulating the Show-Me-State (Lynn 

Robbins, Southwest Missouri State 

University, personal communication). 

Confounding the matter further are the 

controversial predatory and wide-ranging 

characteristics of the animal and local 

uncertainty regarding its actual status as an 

endangered species versus a FERC. Not 

surprisingly then, in the case of the puma in 

the East, Midwest, or prairies where it is


frequently perceived by natural resource 

agencies as a “species non grata”, few 

intensive prospective documentation efforts 

have ever been undertaken (Van Dyke1983, 

McGinnis 1988, Humphreys 1994). To this 

author’s knowledge, east of the Black Hills 

and north of Florida and southern Georgia 

no free-ranging puma has ever been radioinstrumented 

or otherwise marked and 

tracked. 

What we are left with in the case of 

pumas east of the Rockies are largely 

sighting reports and compilations of sighting 

reports. Such data are often met with 

incredulity, yet historically the scientific 

literature, particularly that related to 

carnivores, is replete with papers involving 

nothing more than sighting reports. Articles 

by Berg et al. (1983), involving pumas, and 

Quinn (1995), who worked with urban 

coyotes, are but two of many peer-reviewed 

examples of which this author is aware. 

Where people are reporting unknown or 

unsuspected animals, it often seems one is 

eventually killed and populations are 

subsequently substantiated, vindicating 

those who originally reported sightings. 

Furthermore, the survival of rare animals 

often depends on timely and critical 

decision-making. If sighting reports are the 

best with which a researcher has to work, 

and particularly if some level of scientific 

rigor can be applied to their interpretation, 

as demonstrated by Quinn’s coyote research 

involving sighting reports in Washington, D. 

C., then it is unscientific and ill advised to 

carte blanche ignore such reports (Quinn 

1995). 

In the case of colonization or repopulation, 

by definition there is a temporal 

continuum of occurrence. Early in the 

process, the animals in question are few. 

Colonization, re-colonization, or repopulation 

ends, if successful, with a selfsustaining 

population. The puma 

phenomenon east of the Rocky Mountains is 


presumably somewhere along this 

continuum. Evidence presented at this 

conference suggests puma presence in some 

areas of the West, for instance east-central 

and eastern New Mexico, is on the same 

continuum (Rick Winslow, New Mexico 

Game and Fish Department, personal 


The question of breeding is, of course, 

pivotal in the discussion of purported puma 

populations and presence. Here again, 

however, other than efforts to collect 

sighting reports, there has been little formal, 

proactive modern research on the species in 

eastern North America so information on 

this topic is limited. Nonetheless, some 

useful information can be derived from the 

available data. In the 1970s or early 1980s, 

a string of credible eyewitness reports 

suggested the presence of an adult puma and 

kitten(s) along the Blue Ridge Parkway 

(Robert Downing, USFWS, retired, personal 

communication). A carnivore biologist 

claims to have observed a family group of 

pumas in northern Minnesota in the 1970s 

(Bill Berg, Minnesota Department of 

Natural Resources, retired, personal 

communication). More recently in 

Minnesota, breeding was also implied in the 

case of the female puma killed outside 

Duluth in 2001 (Anonymous 2002). In her 

company were two kittens, both later 

captured and placed into captivity (Mark 

Dowling, Eastern Cougar Network, personal 

communication). The cougar killed in Floyd 

County, Kentucky in 1997, cited earlier, had 

spotted pelage and was in the company of at 

least one other, larger cat, when it was 

struck by a car (Bolgiano 2001). A 

biological scientist observed a puma and 

several kittens in Missouri in the early 

1990s. This is one of several episodes, 

including the poaching of a cougar (Texas 

County, cited earlier) and the videotaping of 

a puma at a deer kill that triggered a 

substantial increase in public and agency

awareness of the puma in the state (Lynn 

Robbins, Southwest Missouri State 

University, personal communication). As 

noted above, recent credible reports of 

females with kittens have also originated 

from both Vermont and Maine. Many other 

instances of apparent puma breeding in the 

East were discussed by Wright (1972). Such 

isolated incidents are certainly not 

unequivocal proof of a puma population or 

breeding, but in the big picture they do tend 

to support the belief that at least a few 

pumas are present and sporadic reproduction 

is occurring. 

Predator Parallels - Bobcat, Black Bear, 

Jaguar, and Coyote 

Many of those skeptical of puma 

presence in eastern North America cite the 

vast suburbanization and urbanization of the 

region as an effective limiting factor. Yet, if 

populations of the versatile puma can exist 

in human dense areas of California, 

Colorado, and Florida, the species could 

surely inhabit portions of the East and 

Midwest, especially given the high 

populations of deer, feral hogs, and other, 

mid-sized and smaller game found 

throughout the region. Furthermore, other 

eastern carnivores including bobcats, black 

bears, and wolves are apparently acclimating 

to evolving habitats and human presence and 

expanding their populations and/or ranges 

(Stoll 1996). Benchmarking with these 

species supports the contention that the even 

more elastic puma can do the same. A 

similar comparison can be made with the 

jaguar (Panthera onca) in the West, which, 

while noted in the region only rarely for 

decades, has been probing borderland 

Mexico-Arizona-New Mexico habitat with 

increasing frequency in recent years (Brown 

and Lopez-Gonzalez 2000). 

The coyote provides an additional case 

study in relation to the possible existence of 

the puma east of the Rocky Mountains 

(Tischendorf 1994b). This adaptable, mid- 



sized, quintessentially western carnivore 

arrived on the midwestern and eastern 

landscape as a veritable newcomer in 

approximately the 1960s and 70s (Gerry 

Parker, Canadian Wildlife Service, retired, 

personal communication). Similar to what 

transpires today with many puma reports, 

coyote presence was initially refuted or 

attributed to feral individuals or 

hybridization with dogs. In retrospect, these 

assessments were not entirely correct. It is a 

convincing reflection of the regional habitat 

quality and prey base that so successfully 

has the species colonized the eastern half of 

the continent it is today a thriving, legally 

trapped, hunted, and pursued game animal. 

If the puma is as adaptable as its history 

suggests, then intuitively it is simply a 

matter of time before it follows the coyote’s 

lead. 

Biolegal Issues 

While current theories support the 

contention that North American pumas are 

genetically panmictic, the Endangered 

Species Act (ESA) specifically identifies 

only the subspecies P. c. couguar (the 

supposed true “eastern puma”) and P. c. 

coryi (the “Florida panther” or more 

correctly “southern puma” [Greenwell 

1996:18, 36]) as endangered (Florida 

Panther Interagency Committee 1993, 

Greenwell 1996, Culver et al 2000). 

Florida, with its mongrel mix of native, 

Texan, “Piper”, and illicitly released 

animals, has overcome this issue by working 

with the federal government to enact 

similarity of appearance protection laws for 

all of its pumas. Thus, Florida’s pumas, 

regardless of origin, now all fall under the 

convenient, albeit taxonomically outdated 

umbrella moniker of “Florida panther” 

(Alvarez 1993). 

Elsewhere in the East and the Midwest 

there exists much confusion and feline 

filibustering about what constitutes a puma 

meriting ESA protection versus one that can


be considered a western wanderer or 

escaped or released captive (Tischendorf 

1994b, Cardoza and Langlois 2002). 

Clouding the issue is the fact that much of 

the Midwest was considered original range 

of F. c. schorgeri, the supposed “Wisconsin 

puma” which technically fits neither into the 

ESA nor the eastern cougar recovery plan of 

1982, which in any case has never been 

implemented (Nowak 1976, Hall 1981, 

USFWS 1982). 

As Albert Einstein reportedly said, 

“Perfection of means and confusion of goals 

seem, in my opinion, to characterize our 

age.” More simply, sometimes the process 

can get in the way of the purpose. Clearly, 

the Endangered Species Act is meant to 

protect rare animals. Equally apparent, the 

puma from the Great Plains eastward is rare. 

Should the ESA unequivocally and 

ultimately serve as a tool to protect this lithe 

and golden ghost as it reestablishes itself 

across its original range? Or can the case be 

made that federal delisting, in concert with 

state or multi-state management plans and 

agreements, unencumbered by federal 

oversight, may more favorably serve the 

puma of the Great Plains, Midwest, and 

East? 

The Jaguar Conservation Team 

(JAGCT), a diverse coalition of agencies 

and individuals working together to develop 

and implement a sound plan for protecting 

and conserving jaguars in the Southwest, 

may serve as a template organization for 

those involved with puma recovery east of 

the Rocky Mountains. Formed in 1997, 

prior to listing of the jaguar as an 

endangered species north of Mexico by the 

USFWS, the JAGCT operates under a 

formal conservation agreement with the 

USFWS and today functions as an ad hoc 

jaguar recovery team (Bill Van Pelt, Arizona 


communication). In essence, the JAGCT 

attempts to preempt the potential for heavy- 


handedness and long-distance directives of 

the ESA by working locally with all its 

stakeholders to proactively find effective 

solutions to conflict (Bill Van Pelt, Arizona 



For now the future of eastern or 

midwestern pumas is largely tied to the 

ESA. Certainly similarity of appearance 

semantics related to pumas and the ESA in 

Florida have symbiotically done much to 

pave the way for puma recovery elsewhere 

east of the Rockies. Still, if the ESA is to 

play a pivotal role, it clearly requires 

modification to recognize P. concolor in 

terms of individuals and populations, 

without regard to an obsolete concept of 

subspecies, as the rare animal that east of the 

Rocky Mountains it truly is (Tischendorf 

1994c, 1995). Such modification would 

thus effectively resolve the unintentional but 

critical double standard for recovery that 

exists for pumas in Florida versus those 

elsewhere in the East and Midwest. Based 

on the powerful southern precedent, 

similarity of appearance protection for all 

free-ranging non-nuisance pumas and 

potential puma populations today and 

tomorrow, not only in Florida, but those 

from the Great Plains eastward, would be a 

simple, logical, and consistent step 

(Tischendorf 1994c, 1995; Cardoza and 

Langlois 2002). 

In the northern Rockies with the wolf, 

and in Florida with the puma, recovery has 

been facilitated by formal restoration efforts. 

It is questionable whether such 

governmentally sanctioned activities will be 

carried out to enhance natural cougar 

recovery in eastern or midwestern North 

America. Economic issues certainly exist, 

witness the budgets necessary for not only 

gray wolf and panther restoration, but those 

for other high-profile endangered species 

like the red wolf (Canis rufus), black-footed 

ferret (Mustela nigripes), whooping crane

(Grus americana), California condor 

(Gymnogyps californianus), and peregrine 

falcon (Falco peregrinus). 

Additionally, given that the species in 

question is not just endangered but large, 

carnivorous, potentially hazardous to 

humans, and an effective ecological 

regulator of ungulates that figure 

prominently in a deeply entrenched hunting 

tradition, there is certainly potential for 

controversy and conflict among various 

public constituencies. 

Still, there are intriguing possibilities for 

the future of the puma in eastern North 

America. Even despite the limitations of the 

ESA, given nothing more than appropriate 

deer management, can the once ubiquitous 

puma survive, re-populate, and thrive in the 

East, Midwest, and Great Plains? Evidence 

presented in this paper tends to support this 

scenario. 

What truly are the public attitudes 

toward this widely ranging and exquisitely 

adaptable carnivore? Do agency attitudes 

mirror public sentiment? These critical 

human dimension wildlife topics would 

make excellent subject matter for a graduate 

student project. 

As pumas return to the Great Plains, 

Midwest, and East, there will inevitably be 

conflict, as occurs in the West, with 

agricultural and suburban interests. 

Exemplifying this, uncannily, as this paper 

was being revised in October 2003, a freeranging 

young male puma, presumably 

dispersing from the west, was captured and 

placed into captivity after causing public 

unrest and alarm in urban Omaha, Nebraska. 

Meanwhile, in Iowa another young male 

puma was shot and killed by a farmer. 

Can we as wildlife professionals devise a 

new ESA or state-level paradigm for 

carnivore management, and specifically the 

phenomenon of novel puma presence, that 

provides effective oversight for scientific, 

evidence-based decisions while allowing for 



sensible managerial flexibility and creativity 

at the key stakeholders’ state and local 

level? Does the JAGCT, with its formal and 

proactive conservation agreement with the 

USFWS, provide a workable model for this? 

Are there in fact several low density, 

highly mobile puma populations and 

breeding foci in eastern and mid-western 

North America? This question remains 

unresolved, but the growing body of 

evidence discussed herein suggests that this 

possibility should not be discounted. 

Finally, despite the substantial evidence 

to the contrary, if pumas are in fact absent 

from the landscapes of the prairies, 

Midwest, and East, what is the prognosis for 

their human-aided restoration in seemingly 

viable ecosystems like the Adirondack 

Mountains, the prey-rich Alleghenies, the 

Ozark or Ouachita Mountains, or the 

expansive northern forests of Minnesota, 

Michigan, and Maine? 

Many of these and other questions were 

raised and addressed in detail at the historic 

“Eastern Cougar Conference, 1994” held in 

Erie, Pennsylvania in June 1994 

(Tischendorf and Ropski 1996). This was 

the first, and remains the only, formal 

conference ever devoted entirely to the 

subject of pumas in eastern North America. 

A second conference is planned for 

Morgantown, West Virginia, in April 2004. 

This conference will roughly mark the tenth 

anniversary of the first gathering, providing 

a centralized forum for ongoing discussions 

and updates on research related to this 

intriguing subject. 

CONCLUSION 

In conclusion, 4 key points: 

1. Adaptable animals adapt and the 

puma is one of the most adaptable 

land mammals in the New World. 

2. There are no ecological reasons why 

the puma could not exist in eastern


North America today. Across much 

of the Great Plains, midwestern, and 

eastern portions of the continent the 

evidence suggests that in low 

densities it does. 

3. Using the model of the JAGCT, a 

diverse but integrated Conservation 

Team should be formed as soon as 

possible to promulgate appropriate 

changes to the ESA (including even 

possible delisting) and to critically 

evaluate the long dormant recovery 

plan for cougars in the East, which 

requires updating to reflect recent 

knowledge related to the puma east of 

the Rocky Mountains (Tischendorf 

1996b, Cardoza and Langlois 2002). 

4. As mankind enters this new 

millennium, return of the puma to its 

former range in the Great Plains, 

Midwest, and East provides an 

opportunity for wildlife professionals 

with limited firsthand experience with 

large carnivores to demonstrate their 

expertise in scientifically and 

sociologically managing this 

relatively secretive predator on the 

many-faceted modern ecological 

interface of private and public lands, 

politics, and public opinion. 

In today’s anthropocentric world, the 

puma, as is the case for large carnivores 

everywhere, is unfortunately a victim of its 

own three “E’s” - its evolution, its ecology, 

and its ethology. Widely ranging, oblivious 

to human-delineated boundaries, a large and 

potentially dangerous predator that preys 

effectively and efficiently on ungulates both 

wild and domestic, the puma is an 

irrefutable, anachronistic, and controversial 

symbol of our primeval wild. 

In the end, the message that emerges 

today for the puma in the East is this: even 

at its highest densities, few people will ever 

see a living, wild puma. Certainly we can 


live without this great cat. Even more 

certainly, it can live without us. Enmeshed 

in controversy, entrenched in folklore, 

history, and legend, endangered across a 

huge portion of its historic range, the largely 

secretive puma presents us with the great 

challenge of, and the magnificent 

opportunity for, harmonious coexistence. 

Hopefully mankind will rise to this rare 

occasion to ensure that the puma is again a 

celebrated and wisely managed part of our 

Great Plains, midwestern, and eastern 

wildlife heritage. 

ACKNOWLEDGEMENTS 

This paper is dedicated to the memory of 

Frank C. Craighead, Jr., of Moose, 

Wyoming, whose life and career were an 

inspiration to a generation of wildlife 

biologists. Kerry Murphy and Randy 

Matchett served knowledgeably and capably 

as manuscript referees. A special thank you 

is extended to the Wyoming Game and Fish 

Department, coordinator and host for this 

Mountain Lion Workshop. 


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MOUNTAIN LION STATUS REPORT: BRITISH COLUMBIA 

MATT AUSTIN, Large Carnivore Specialist, Ministry of Water, Land and Air Protection, PO 

Box STN PROV GOVT, Victoria BC V8W 9M4, Canada, email: 

Matt.Austin@gems7.gov.bc.ca 

Abstract: Mountain lions are classified as “Big Game” in British Columbia under the provincial 

Wildlife Act. There is no provincial mountain lion management plan, however, there is a species 

account within the provincial Wildlife Harvest Strategy. The harvest management goal for 

mountain lions is “to optimize population sustainability within ecosystems while allowing for 

options and opportunities associated with hunting and viewing.” Mountain lions occupy all 

suitable habitats within BC. The distribution of mountain lions has been expanding northward in 

recent years due deer population increases resulting from less severe winters. The current 

provincial mountain lion population estimate is 4,000-6,000 and likely declining after peaking in 

the late 1990s. Declines are related to the severe winter in 1996/97 that reduced deer 

populations. Population estimates are based on the “best guesses” of regional biologists based 

on anecdotal and harvest/conflict information. Confidence in the population estimate is low but 

we have greater confidence in the trend estimate. Mountain lion hunting is allowed under 

General Open Seasons in all but two northern regions with negligible populations. There are no 

explicit harvest criteria or objectives aside from quotas for female harvest in one region. Both 

harvest and mortalities resulting from conflicts increased from 1985 until 1996 and then declined 

through 2002. Conservation Officers respond to conflicts with mountain lions through 

education, translocation or destruction; compensation is not provided for losses. Known 

mountain lion attacks increased during the 20 th century, peaking in the 1990s. 


87

88 

IMPROVING OUR UNDERSTANDING OF MOUNTAIN LION MANAGEMENT 

TRENDS: THE VALUE OF CONSISTENT MULT-STATE RECORD KEEPING 

CHRISTOPHER M. PAPOUCHIS, Mountain Lion Foundation, PO Box 1896, Sacramento, CA 

95814, USA, email: cpapouchis@mountainlion.org 

LYNN MICHELLE CULLENS, Mountain Lion Foundation, PO Box 1896, Sacramento, CA 

95814, USA, email: cullens@mountainlion.org 

Abstract: The sound management and conservation of mountain lions relies on comprehensive 

scientific data. Yet the cost of mountain lion research can be prohibitive and the results are often 

difficult if not impossible to extrapolate. Wildlife managers, field researchers, and conservation 

organizations would benefit from more complete and consistent records of validated mountain 

lion sightings, hunting mortalities, depredation incidents, and road kills. Scientists who have 

mined such data in the past have isolated important variables, generated important hypotheses, 

and targeted future research. But their work is usually limited by funding, academic or agency 

agendas. Further, there is no long-term multi-state repository for mountain lion data. The task 

of data collection is made more difficult because there is no multi-state standard, and therefore 

states collect and store data inconsistently. This presentation explores the potential for 

developing a multi-state database, and examines the existing state data sets in order to identify 

the essential variables that might be included. 

PROCEEDINGS OF THE SEVENTH MOUNTAIN LION WORKSHOP

LESSENING THE IMPACT OF A PUMA ATTACK ON A HUMAN 

E. LEE FITZHUGH, Department of Wildlife, Fish, and Conservation Biology, University of 

California, Davis, One Shields Avenue, Davis, CA 95616-8751, USA, email: 

elfitzhugh@ucdavis.edu 

SABINE SCHMID-HOLMES, Department of Wildlife, Fish, and Conservation Biology, 

University of California, Davis, One Shields Avenue, Davis, CA 95616-8751, USA 

MARC W. KENYON, Department of Wildlife, Fish, and Conservation Biology, University of 

California, Davis, One Shields Avenue, Davis, CA 95616-8751, USA 

KATHY ETLING, 6830 St. Tropez Circle, Osage Beach, MO 65065, USA 

Abstract: We reviewed current data on puma (Puma concolor) attacks and near-attacks on humans to identify better 

ways for people to protect themselves. Not since Paul Beier’s paper in 1991 has anyone documented, established 

criteria for validity, and analyzed puma attacks on humans, and much more data are now available. In attempting to 

examine human-puma behavioral interactions to 2003, the authors have collected accounts of 16 fatal and 92 nonfatal 

attacks that meet Beier’s criteria. In addition, we have an additional 32 fatal and 84 non-fatal attacks that failed 

to meet Beier’s criteria, either for lack of physical contact, lack of verification, occurrence in Latin America, 

occurrence prior to 1890, or because they were attacks on hunters. We also have accumulated 155 accounts of 

behavioral interactions between pumas and humans at close proximity that did not result in an attack. We contrasted 

these with incidents that resulted in an attack. We analyzed the use of Beier’s fatal:non-fatal attack ratio to predict 

missing incidents, and suspect that the criterion of validation may bias data for attacks prior to 1950. However, 

most of Beier’s statements and conclusions are confirmed. While the analysis is yet incomplete, this presentation 

includes highlights of our tentative analysis concerning common questions about puma attacks, illustrated by stories 

of real situations. Being aggressive and making loud noises helps protect people from a possible puma attack. 

Warning gunshots are much less effective than is yelling. Charging the puma seems to make it run away, but may 

result in some injury to the person who is charging. Groups of 5 people or more are relatively safe, but children in 

those groups may still be attacked. Hunters imitating animal sounds or smells may attract pumas, but these 

situations usually do not result in serious injuries. People attacked while sleeping on the ground often receive only 

minor injuries because the puma runs away when the person or companions awake, yell, and resist. The strategies 

will usually work, but not always, because pumas have different personalities and seem to react differently to the 

same situation. 

89 


Key words: Animal damage, attacks on humans, conflict with wildlife, cougar, human dimensions of wildlife 

management, mountain lion, pest control, predation, Puma concolor 

INTRODUCTION 

In this paper we use the common name 

“puma” to describe Puma concolor. 

Occasionally, when we quote other people, 

we retain their terminology, and they often 

use “cougar” instead of “puma.” Beier 

(1991) analyzed 9 fatal and 44 non-fatal 

attacks by pumas on humans that occurred 

between 1890 and 1990 in the United States 

and Canada. In order to include an attack in 

his analysis, it must have been published, 

included statements from agency or medical 

personnel, and involved contact in which the 

human was bitten, clawed, or knocked down 

by the puma. Excluded were situations 

involving captive pumas and in which 

people deliberately approached or harassed a 

puma. He found that 64% of victims were 

children, and only 13 of 37 (35%) of these 

were alone, while 11 of 17 (65%) adult 

victims were alone when attacked. He also 

found that an aggressive response might 

avert and/or repel an attack. Yearlings and 

underweight cougars were most likely to

90 REDUCING PUMA ATTACKS · Fitzhugh et al. 

attack humans. Beier believed he had 

discovered all fatal attacks since 1890 that 

met his criteria, and all non-fatal attacks 

since 1970. Based on the ratio of fatal to 

non-fatal attacks during 1970-1990, he 

estimated that he had failed to identify about 

12 non-fatal attacks between 1890 and 1970. 

Beier documented an increase in frequency 

of attacks from the 1890-1969 period to 

1970-1990. While Beier did not tabulate 

“near-attacks,” he did analyze the victim’s 

actions that may have served to prevent the 

attack. “Fighting back” and shouting loudly 

were actions that seemed to avert or repel 

attacks, as did waving arms, poking or 

hitting with sticks, throwing rocks, etc. 

Beier also reported an attempt at aversive 

conditioning of one puma, but it failed to 

prevent future aggression. We substantiated 

most of Beier’s findings and, because we 

have more data, we produced some 

additional tentative findings. 

We have accounts of a total of 224 

attacks by pumas on humans and 155 

behavioral interactions that did not result in 

an actual attack. The number of accounts 

through April 2003 that have information 

useful for analyzing any specific question is 

variable, but only 108 accounts meet Beier’s 

(1991) criteria. Of the 116 attacks that failed 

Beier’s criteria, 32 were fatal and 84 were 

non-fatal. Reasons for failure include lack of 

physical contact, lack of verification, 

occurrence in Latin America, occurrence 

prior to 1890, or because they were attacks 

on hunters. Beier’s strict criteria avoid 

errors of commission, but allow for many 

omissions. Because we are interested in 

behavior more than just counting attacks, we 

decided to relax the Beier limitations as long 

as accounts seemed plausible and contained 

useful information. Our intent is to analyze 

human and puma behavior in these 

situations where data are available and 

compare attacks with encounters to try to 

provide better advice for people who come 


face-to-face with a puma, or who want to 

prepare for that eventuality. We are still 

organizing the data, so this report is not the 

final one, but we do have a few 

recommendations to make at this time, and 

we will make some observations about 

reliability of reports and frequency of 

attacks in general. 

METHODS 

We defined “non-attack encounters” as 

behavioral interactions between pumas and 

humans at close proximity that do not result 

in an attack. We purposely did not define 

“close proximity” to place emphasis on 

“behavioral interactions.” We excluded 

incidents in which the puma was sighted and 

then left, and included incidents in which the 

puma and human exchanged multiple 

behaviors. “Close proximity” is necessary 

for this to occur, but the distance may vary, 

and we were more flexible regarding 

distance criteria than for behavioral criteria. 

If we believed we could learn from the 

interaction, we included it. Most of our data 

are from published popular accounts, 

sometimes substantiated by an agency 

incident report. Etling (2001), in particular, 

solicited personal accounts from individuals 

in their own words, both before and after 

publication of her book. We categorized 

incidents in several ways to better analyze 

and evaluate the data. One mentioned in 

this paper is a category we called “attacks 

terminated by humans.” These are incidents 

in which a puma was shot while charging, or 

at least clearly intent on creeping up very 

close to a human in spite of the person’s 

efforts to discourage such behavior. We 

have 20 such accounts, 10 of which are from 

hunters. In only 3 of the hunting accounts 

was it clear that the hunter was doing 

something that might attract a puma (e.g., 

using deer scent, calling turkeys, etc.). Six 

accounts were of agency employees 

investigating previous encounters between 

humans and pumas.

We defined a child as a person under 13 

years of age, whereas Beier (1991) defined a 

child as being under 16. We differed from 

Beier because we believed that the younger 

age better represented when girls and boys 

reach adult size and behavior. 

We entered data into a spreadsheet and 

made inferences where they were defensible. 

For example, when the victim fired a shot, 

and the attack was fatal to the victim, we 

inferred that the shot did not deter the 

attacking puma, even though the account did 

not specifically say so. The matrix was 

organized with individual incidents in rows, 

separated into various categories such as: 

Beier fatal, Beier non-fatal, non-verified 

(otherwise meeting Beier’s criteria), fatal 

prior to 1890, nonfatal prior to 1890, Latin 

American incidents, close encounters, 

provoked attacks, encounters while hunting, 

and 10 other categories. Columns included 

raw data and data coded into categories for 

analysis, all of which made 193 columns, 

including 147 columns of original data. 

Broad categories of data in columns 

included descriptions of the habitat and 

setting, identification and descriptions of the 

victims, and detailed descriptions of the 

incident, including a written description and 

data parsed into separate columns. 

Information sources, previous reported 

puma activities in the area, necropsy results, 

and injuries sustained by the victim(s) also 

were entered. An example of data sought 

and entered is in the partial questionnaire 

mentioned below. 

On a few recent occasions, when it was 

possible, we sent a 4-page questionnaire to 

the witness of an attack to get more detailed 

information. The questionnaire was generic 

and designed to help people remember detail 

without leading them to specific answers. 

Thus, we did not ask whether the puma was 

pumping its rear feet up and down, but 

instead we asked what the puma was doing 

with its feet. Although the situations were 


REDUCING PUMA ATTACKS · Fitzhugh et al. 91 

serious, the responses sometimes interacted 

with the generic nature of the questions in a 

humorous manner. A real example of a 

questionnaire we recently received from a 

man who gave permission to use it will 

illustrate both the detail of the questions and 

some of the humor that occasionally occurs. 

It also illustrates an incident that is not 

classified as an attack under the Beier 

(1991) criteria, because no contact was 

made. We have called it a “terminated 

attack.” 

To appreciate the humor, it helps to 

imagine the victim’s perspective on the 

attack and his response to the “ivory-tower” 

nature of the questions that came from some 

stranger at a far-off university. The victim 

was hunting deer in a remote area when he 

was charged from behind. The puma 

vocalized with a “growl-hiss” sound, which 

alerted the victim to the charge. He killed 

the puma during its charge, after missing his 

first shot. After the second shot the puma 

fell only 5.2 m (17 feet) from him. Here are 

a few of the questions and the victim’s 

responses: 

How was the puma identified (what 

evidence or characteristics)? 

The cougar died, not much question that 

it was a cougar. 

Condition of teeth: 

Perfect teeth, no fillings. 

Condition of claws: 

Damned sharp. 

Did attack involve a fatality? 

Yes, the cougar. 

Puma posture and position of ears at time of 

first sighting: 

The cat was charging me. I later 

measured the distance from where it 

started the charge, which was 86 feet. I 

don’t recall what position the ears were 

in. [86 feet is 26.2 m]. 

What was puma doing with eyes and tail at 

time of first sighting? 

Tail seemed to be floating out behind the


cat. 

What was puma doing with its feet at time 

of first sighting? 

Bounding toward me. 

Victim behavior just after first sighting? 

Putting rifle to shoulder and firing. 

Were there signs of aggression by puma? 

The cougar was charging me, full speed 

ahead, which seemed pretty aggressive to 

me at the time. 

Did victim fight back? 

Yes. 

How? 

I shot the puma in the throat/chest. 

Puma response: 

Puma rolled and died. 

Was puma injured by victim? 

Yes, severely. 

Quality of Data 

Some accounts are not included in this 

analysis. The 224 attacks and 155 

encounters we analyze do not include 14 

incidents for which we believe additional 

investigation is needed to validate their 

accuracy. Nor do we include 8 other 

incidents we suspect, but cannot prove, are 

duplicates of incidents included in the tally. 

In addition, we have 10 more reports that we 

decided not to use because they included too 

little information or were of doubtful 

validity. Our data do include incidents that 

do not meet Beier’s criteria, but we kept 

those separate in order not to invalidate 

comparison with Beier’s (1991) findings. As 

we analyze data more completely, more of 

the incidents may be excluded, primarily for 

lack of information. Our files also include 

several accounts that have recently come to 

our attention that are not yet entered in the 

database and are not included in this paper. 

These latter data, and additional details from 

the accounts we have analyzed will be 

included in a later, more complete treatment. 

As already mentioned, we included all 

the accounts we have discovered, and we 

tried to estimate the validity of each. The 


editor of Outdoor Life, from 1897 to 1925, 

asked his readers to provide accounts of 

puma attacks on humans. He then tried to 

verify the accuracy of each account, 

generally without success. In each case, he 

was either unable to locate the respondent, 

or the knowledgeable people from the area 

where the attack was supposed to have 

occurred claimed no knowledge of it 

(Anonymous 1925, cited by Beier 1991). 

During this same period, Forest and Stream, 

which became Field and Stream, also 

printed numerous personal accounts of 

encounters with pumas. At this time, we 

have been able to locate only 1 reference to 

incidents that may have been confirmed 

(Marsh 1917), or failed confirmation, by 

Outdoor Life (Anonymous 1917). We are 

aware of one, and perhaps three fraudulent 

accounts in recent years, and we also 

questioned the validity of one unusual 

account that we later found had been 

confirmed by an agency. We recently tried, 

unsuccessfully, to obtain agency 

confirmation of an account, only later to find 

that the confirmation had been provided to 

Etling several years earlier. Therefore, the 

verifications themselves can be erroneous in 

either direction. It is possible that we have 

analyzed a few spurious reports, but if so, 

their effect on our findings should be minor. 

We have placed our 379 useful incidents 

in categories of similar types of incidents 

and levels of reliability. The 108 fatal and 

non-fatal incidents that meet Beier’s (1991) 

criteria may be considered to be a complete 

count of well-defined and verified attacks. 

(See a more complete defense of this 

assumption in the results and discussion 

section). The few verified non-fatal 

incidents we may have missed would not 

affect group values in an important way. 

The 116 other attacks and 155 encounters 

represent neither a total count nor a 

statistical sample, nor do we know anything 

about the underlying statistical distribution

except that attacks by pumas on humans are 

rare. We know nothing of bias caused by 

missing data. We can speculate that missing 

cases may not have been considered 

important enough to report; there may have 

been nobody to report to; fatal incidents may 

have been undiscovered; they may have 

been reported and the records lost; they may 

have been printed in obscure references that 

we did not find; we may have wrongly 

discarded some incidents recorded from 

word-of-mouth accounts, etc. Thus, we 

have a core of strictly defined data that we 

treat as a total count. These data are 

restricted by the verification criterion in 

such a way that the passage of time reduces 

opportunity for verification. The core exists 

in a matrix of less well-defined incidents, 

the statistical properties of which are 

unknown. 

At a finer level, even the welldocumented 

cases have many blank cells in 

the data matrix because specific items were 

unknown or not reported, and there was no 

way to infer the information. In these 

situations, we usually cannot assume the 

nature of possible biases caused by missing 

data, if they exist. 

To summarize, we have nearly total 

counts of incidents in Beier-quality 

categories, missing incidents in others, and 

missing data in all incidents. The categories 

in which we can assume total counts may be 

subject to a time-related bias that may affect 

the statistical distribution of the data. We 

have little a priori information to guide us. 

Therefore, we are restricted mostly to 

descriptive statistics and forming hypotheses 

that may later be independently verified. 

We do use a Chi-square test to explore the 

similarity of the distributions in three 

different categories, two of which we 

assume are total counts. 

Statistical Methods 

We have no a priori models or 

hypotheses. We had believed that most, if 



not all, puma attacks were predatory, but 

information provided by Sweanor et al. 

(personal communication) contradicts that 

belief. Thus, our analysis is exploratory, 

examining the data to find hypotheses and 

descriptive models that may later be 

subjected to data collection and statistical 

interpretation. In the few cases where 

statistical testing was warranted, we report 

the test used along with the results, but for 

the majority of situations, descriptive 

statistics are the only analysis used. 

Nevertheless, we feel secure in drawing 

some conclusions about how to reduce risk 

of serious injury during a puma encounter. 

RESULTS AND DISCUSSION 

What Can We Tell About the Data? 

With respect to counting attacks and 

comparing frequencies, we can be a little 

more specific about statistical qualities of 

the data. Like Beier (1991), we believe we 

have a near-complete count of verified fatal 

attacks from 1890 to 2003 in the U. S. and 

Canada. Authors of new books (Danz, 

1999, Deurbrouck and Miller 2001, and 

Etling 2001) did extensive new searches, 

and failed to find any fatal attacks that meet 

Beier’s criteria that were not included by 

Beier in his original list, or else occurred 

after his publication. The 108 attacks we 

analyze that meet Beier’s criteria include 7 

fatal and 38 non-fatal attacks that occurred 

after Beier published his list, and 9 fatal and 

54 non-fatal attacks that meet Beier’s 

criteria and occurred between 1890 and 

1991. On the basis of additional information 

(personal communication, Dale Elliot to 

Etling, July 2000), we moved one of Beier’s 

non-fatal attacks (Bird and Sieh, Nevada, 

1971) to the “provoked attack” category and 

added 11 new non-fatal attacks between 

1890 and 1991. It is possible that attacks, 

even fatal ones, occurred in the U. S. that 

were never known, especially during the 

depression years of the 1930s and the 

various gold rushes in localities in the


Table 1. Calculations of non-fatal attacks by pumas in the U.S. and Canada between 1890 and 1969 

that might not have been detected. 

Source 

Beier (1991) 

1970-1990 

Our data 

1970-2001 

Ratio NF/F 

from 1970 

onward 

× Fatal 

attacks 1890- 

1969 

= Calculated 

non-fatal 

attacks 

− Attacks 

detected 

1890-1969 


= Calculated 

attacks not 

detected 

31÷5 = 6.2 4 25 13 12 

69÷12 = 5.75 4 23 21 2 

western U.S. It is certain that attacks 

occurred in Latin America for which no 

records are available. Written accounts of 

attacks occur in some obscure publications, 

not susceptible to easy location. One 

example is the killing of Henry Ramsey in 

1876 (Hunter 1922:110). That account was 

found by scanning the table of contents of a 

county history that was in a fund-raising 

auction of “white elephant” donations. All 

of these missing accounts would, of course, 

fail Beier’s criterion of verification, and 

perhaps other criteria as well. We believe 

we can use the 16 fatal attacks that meet 

Beier’s criteria as a complete count, not 

requiring statistical measures of variability. 

The 92 Beier-quality non-fatal attacks 

probably include a large majority of all nonfatal 

attacks (Table 1). They should be 

representative, and probably can be treated 

as a complete count. However, they were 

not sampled according to a statistical design. 

The remaining 116 attacks and 155 

encounters have unknown statistical 

properties, with variable report quality and 

amounts of information. 

Beier (1991) estimated that he might 

have missed finding 12 non-fatal accounts 

between 1970 and 1990. He did this by 

assuming that he found all the accounts from 

1970 to 1990, and multiplying the ratio of 

non-fatal to fatal attacks during that period 

by the number of fatal attacks from 1890 to 

1969. We discovered 8 of the 12 missing 

accounts, and also 3 more between 1970 and 

1990, so we recalculated the potentially 

missing non-fatal accounts (Table 1). These 

were calculated through 2001, as the 2002 

data are yet incomplete. The analysis in 

Table 1 assumes a constant rate of attacks 

across years. Puma populations, prey 

populations, and the number, age, sex, and 

group size of people at risk may have 

changed considerably since 1890. These 

factors may affect the attack rate. Thus, the 

calculation may be invalid to the degree that 

these parameters have changed. 

Even if the analysis in Table 1 is valid, 

the number of fatal attacks is small enough 

that a change in even one attack can alter the 

calculation of non-detected non-fatal 

attacks. Because there is a chance that we 

may have missed some fatal attacks prior to 

1970 (and especially prior to 1950 as 

discussed later), this is a tentative 

calculation that serves only to illustrate that 

there likely are some incidents we have not 

found, but that number is relatively small. 

We will return to this topic later with respect 

to possible bias caused by Beier’s 

verification criterion. 

Comparing Verified and Unverified Data 

Figure 1 shows the relationship through 

time between the 15 Beier-quality fatal 

attacks, the 86 non-fatal attacks, and the 27 

non-verified, non-fatal attacks, 1890-1999. 

The Beier-quality non-fatal attack curve

Figure 1. A comparison of patterns of fatal 

and non-fatal attacks that conform to Beier’s 

(1991) criteria with non-verified non-fatal 

attacks. 

diverges from the other two beginning in 

1950. All three types of data have been 

subject to the same bias from a conscious 

increase in collecting attack reports 

beginning with Barnes (1960), our effort 

from 1984 (Fitzhugh and Gorenzel 1986), 

and intensive searches beginning about 1990 

(Beier 1991) and increasing in 1998-2001 

(Danz 1999, Deuerbrouck and Miller 2001, 

Etling 2001).The Beier-quality fatal attacks 

curve (Figure 1) began to exceed past levels 

in the 1970s, and increased even more in the 

1990s. We confirmed that Beier 

documented all the verified fatal attacks 

since 1890, and Figure 1 shows that the nonverified, 

non-fatal attacks coincide closely 

with Beier-quality fatal attacks. The 

difference in the shape of the curves in 

Figure 1 may be partly attributable to an 

increase in agency funding, staffing, and 

attention to puma incidents beginning in 

about 1950, allowing for more verification 

and recording of non-fatal incidents (Harley 

Shaw, personal communication). Thus, the 

post-1950 non-verified data may be 

depressed because a greater proportion of 

those incidents were verified than happened 

pre-1950. The post-1950 verified non-fatal 



Figure 2. Non-fatal to fatal attack ratios. 

Bars represent a 20-year running average 

beginning with 1890-1909, 1900-1919, etc., 

except that 1990 represents only 1990-1999. 

Includes only data that conform to Beier’s 

(1991) criteria. The zero value at 1890 

represents a 20-year period, 1890-1909, with 

no verified non-fatal attacks. The zero value 

at 1950 represents a 20-year period, 1950- 

1969, without fatal attacks. (N = 101; 15 = 

fatal, 86 = non-fatal.) 

data would have increased by the same 

amount. 

The proportional change in the Beierquality 

non-fatal attack curve after 1949, 

applied to higher numbers of non-fatal 

incidents compared with fatal incidents, 

magnifies the visual comparison between 

the curves, although prior to 1950 the nonfatal 

curve is only slightly higher than the 

fatal curve. The effect of magnification can 

be removed by examining proportions 

directly, using non-fatal to fatal ratios by 20year 

periods (Figure 2). The 1960-1990 

ratios seem consistent, with an average 

value of 6.5. The 1890-1930 average is less, 

at 2.4 (including 1890-1909, when there 

were no recorded non-fatal attacks), but is 

more variable. Only the 1940-1959 ratio 

seems unusually high, created by the first 

big increase in non-fatal attacks, which 

occurred during the 1950s, while no fatal 

attacks occurred 1950-1959. While non-


Table 2. The effect of including non-verified puma attack incidents on the non-fatal:fatal attack 

ratios before and after 1950, U. S. and Canada. 

Data 1890-1949 1950-1999 Difference 

Beier-quality only 6 ÷ 4 = 1.5 86 ÷ 12 = 7.2 5.7 

Beier-quality & all non-verified 16 ÷ 7 = 2.3 96 ÷ 13 = 7.3 5.0 

Beier-quality & non-verified non-fatal only 16 ÷ 4 = 4.0 96 ÷ 12 = 8.0 4.0 

fatal attacks did not decrease after that, fatal 

attacks increased starting in 1970, bringing 

the ratios down. Some of the variation in 

Figure 2 is caused by zero values. The nonverified, 

non-fatal data (Figure 1) are 

approximately of the same value as the 

Beier-quality fatal data, and the curves are 

very similar, so we may be justified in 

combining the non-verified non-fatal data (n 

= 23) with the Beier-quality non- fatal data 

(n = 86). Four non-verified fatal attacks also 

were included with the 15 verified fatal 

attacks to be consistent. Ratios for the 

periods before and after 1950 (Table 2) 

show that adding all non-verified incidents 

(fatal and non-fatal) increased the ratios 

slightly; ratios were even greater when only 

non-verified, non fatal incidents were added. 

The non-verified data also reduced the 

differences between the earlier and later 

periods. If we assume that the underlying 

ratio of non-fatal to fatal attacks is 

consistent across years, it appears that 

excluding non-verified data changes the 

ratios. The changes represent bias if we are 

justified in using the non-verified data. 

Beier’s (1991) calculation of ratios from 

1970-1990, to estimate missing non-fatal 

attacks prior to 1970, assumed a constant 

relationship. It seems logical that we have 

detected a larger proportion of the actual 

non-fatal attacks in recent years, and the 

data seem to indicate that this is so (Figures 

1-3). However, some biological, 

demographic, and cultural differences may 

have caused a change in ratios not related to 


reporting frequency. These changes have to 

do with changes in persecution of pumas, 

especially before, during, and after World 

Wars I and II, the number of people using 

puma habitat, changes in the degree of puma 

habituation to humans, changes in the 

proportion of children versus adults exposed 

to pumas, and changes in the inclination or 

ability of people to report incidents. We 

believe the underlying non-fatal to fatal ratio 

may have changed about the middle of the 

20 th century, but measurement of this 

potential change is confused by changes in 

Figure 3. Non-fatal to fatal attack ratios. 

Bars represent a 20-year running average 

beginning with 1890-1909, 1900-1919, etc., 

except that 1990 represents only 1990-1999. 

Included are data that conform to Beier’s 

(1991) criteria and non-verified data, both 

fatal and non-fatal. The zero value at 1950 

represents a 20-year period, 1950-1969, 

without fatal attacks. (n = 128; 19 = fatal, 109 

= non-fatal.)

the rate of detection of attacks and changes 

in verification of detected attacks. 

We did subject the Beier-quality data to 

2 Chi-square comparisons with 27 of the 

accounts that were not verified, but met 

other Beier criteria. These are the same data 

shown in Figure 1. We used the Beierquality 

data as the observed value and the 

non-verified non-fatal data, paired with the 

Beier-quality data by decades, as the 

expected value. The Beier- quality fatal data 

were not different from the non-verified, 

non-fatal data (χ 2 = 13.7, 10 df, P =


Figure 4. Puma responses to noise, including 

lethal shots from firearms (n = 133). 

to draw out the duration of the sound. For 

example, a puma was recently stalking 

chickens in the yard of a lighthouse 

compound, with people around and active, 

in broad daylight. One man banged doors, 

without effect, then got a .22 rifle and fired a 

shot into the ground without even causing 

the puma to flinch or look up. It was fixated 

on the chickens. The man then fired 7-8 

shots rapidly into the ground, upon which 

the puma looked up and walked into the 

nearby brush, but did not leave the area. It 

stayed on a nearby high area and watched 

while people put the chickens into a pen 

(Robert Hansen, Pacific Rim National Park, 

Vancouver Island, B.C., personal 

communication, 8 May, 2003). This 

indicates that the puma did not react to a 

single shot or to slamming a door, but did 

react to a subsequent rapid series of shots. 

We conclude that noise is effective, but 

the kind of noise makes a difference. The 

best deterrent in the event of a puma 

encounter is to yell or scream as loudly as 

possible. If you are going to shoot a gun, 

you should fire in rapid succession to 

frighten the animal away or shoot to kill the 

puma. 


What if You Charge the Puma? 

Our data include 6 accounts in which the 

primary victim either charged the puma and 

fought with it, or engaged in “mock lunges” 

toward the puma. In 3 cases in which 

people actually charged and made contact 

with the puma, the puma left the area, 

sometimes after a brief scuffle in which the 

human suffered light degrees of injury. Two 

examples follow: a man heard a commotion 

in his back yard and went to investigate. He 

thought his Scottie dog was being attacked 

by a large German shepherd. It was really a 

mountain lion, but he didn’t realize it until 

he had jumped onto the attackers back. 

When the man realized it was a puma, he let 

go after a brief scuffle and the puma ran off. 

The man received stitches for cuts behind 

his ear (Colorado Division of Wildlife 

2002). In the other case, a man noticed a 

puma eating his daughter’s house cat, and 

decided to save the cat by wrestling with the 

puma. The puma swatted the man in the 

face, and the man then decided to let go. The 

puma left with the house cat in its mouth 

(The New York Times 2002). 

Two cases involved repeated “mock 

lunges” by people causing the pumas to 

leave the area without attacking. In the first 

case, a woman came upon a puma crouched 

about 1.8 m (6 feet) away. It began to move 

toward her in a crouched position, growling. 

She lunged forward, holding arms wide and 

growled back at it. It retreated a bit, began 

to approach again. She growled and lunged 

again; the puma retreated again, not as 

startled as it was the first time. Then the 

puma took a last glance and turned into the 

forest. The woman walked backwards 

awhile, then turned around and ran (Personal 

correspondence to K. Etling on 4 Dec, 2001, 

from K. Hogland). 

In the second example, a puma 

confronted 2 biology students gathering data 

in Alum Rock Park, San Jose, California. 

The women yelled and made themselves

look bigger, but the cat continued to 

advance. One of the women snarled like a 

dog and “mock lunged,” and the puma ran 

into some bushes. A nearby rancher 

approached on horseback, accompanied by 2 

dogs. When they directed their attention 

toward where the girls thought the puma 

was, it bounded off (Linda Lewis, web site: 

, citing personal communications 

with Jessie Dickson, April 18-19, 2001). 

In probably the most dramatic example 

demonstrating puma behavior following 

human aggressiveness, a deer hunter and a 

puma were stalking the same deer when the 

deer detected the puma and fled. From 27 

m (30 yards) away, the puma transferred its 

stalk to the hunter. The hunter hid behind a 

tree while the puma approached, crouching. 

As the puma got close the hunter jumped out 

and yelled. That puma left running (Ford 

1994). The puma obviously knew the hunter 

was behind the tree, but the hunter’s actions 

probably appeared to the puma as an attack 

coming from a hidden (ambush) position. 

The action successfully interrupted the 

predatory stalking behavior and instigated a 

flight behavior. 

Is It Safer to Hike in Groups? 

Solitary people are 3 times as likely to 

be attacked or to have an encounter as 

people in pairs or larger groups (Figure 5). 

However, only groups of 5 or more seem 

fairly secure against attack. We were much 

less likely to find data on non-attack 

encounters than on attacks. Thus, the 

relative levels of the paired bars in Figure 5 

cannot be used to compare attack and nonattack 

encounters. We assumed that the 

reporting rates for attacks and non-attacks 

are different but are not affected differently 

by group size. Figure 5 shows that the 

relationship (but not absolute proportions) of 

attack and non-attack encounters is similar, 

regardless of group size. The similar 

percentages within group size categories 



Figure 5. Relationship of human group size 

and age composition with type of encounter 

(n = 379). 

and the consistent pattern among them 

indicates to us that the tendency of a puma 

to approach humans (or for humans to come 

close to pumas) is related to group size and 

is independent of whether an attack occurs. 

It seems to indicate that once a puma is in 

close proximity to humans, whether an 

attack occurs or not may be explained, 

statistically speaking, as a random or 

systematic decision, affecting all group sizes 

to the same extent once the initial approach 

is made. Such a mechanism could be 

created either by the 

physiological/behavioral state of the puma 

or size and behavior of the human(s), or 

both interacting. If we could detect and 

record non-attack encounters as thoroughly 

as we do attack encounters, we might be 

able to create more hypotheses based on the 

ratios and timing of one to the other. Data 

presented by Sweanor et al., at the Seventh 

Mountain Lion Workshop, help 

considerably in this direction. We 

encourage all who study radio collared 

pumas to record and publish similar data. 

In Figure 5, adults strongly predominate 

in the single person attack and non-attack


Figure 6. Proportions of human age classes 

in different types of incidents, as affected by 

human group size (n = 379). 

categories and in the two-person non-attack 

category. This may reflect the relative use 

of wildland trails by single and paired adults 

versus children. However, the categories of 

attacks on two people and on 3-5 people 

show increasing proportions of groups 

mostly composed of children, while the nonattack 

categories for the same size groups 

still show a predominance of groups mostly 

composed of adults. This indicates that, 

while attacks on groups of two or more 

people are much fewer than attacks on 

individuals, children in these larger groups 

are at only slightly less risk than children 

alone. 

Another way of viewing the same data is 

to scale each column separately and 

proportion the age groups within columns 

rather than being based on total incidents 

(Figure 6). When we do this, the increased 

proportion of attacks on mostly-children 

groups in the larger group sizes becomes 

more noticeable, although sample size is 

small (n = 16 for children in groups of >2 

people; n = 17 for adults in the same size 

groups). Among the 17 groups of 3 or more 

people composed mostly of adults, children 

were attacked in 11 of those groups. For 


example, a group of 2 or 3 children (11-12 

years of age) plus 5-6 adults were on a 

kayak tour, camped on Compton Island in 

Johnstone Strait, V.I., B.C. recently. The 

group had just gotten out of the kayaks and 

was on the beach, standing in a group with 

the children in the middle, when a 100-lb. 

puma charged out of the brush, grabbed a 

12-year-girl and dragged her off. Shouting 

and noise made by the adults made the cat 

drop the girl, who survived the incident 

(Robert Hansen, Pacific Rim National Park, 

Vancouver Island, B.C., personal 

communication, 8 May, 2003). 

Is It Safe to Sleep on the Ground? 

We know of 12 victims who were 

accosted in their sleep (Figure 7). Only one, 

a man in Argentina in 1898, was seriously 

injured, and details of that account are 

lacking (Pritchard 1902; Roosevelt 1914:29; 

Barnes 1960:119,122). Seven victims were 

uninjured, and 4 others suffered minor 

injuries. For example, a boy was sleeping 

on a mat at night when a puma walked up, 

pawed him, and tried to drag the mat away. 

His father rescued him. The boy suffered 

scratches and had 10 stitches in his ear. 

(Phoenix Gazette and Tucson Citizen 1994; 

Danz 1999: 276-277; personal 

Figure 7. Injuries that occurred when victim 

was attacked while sleeping (n = 12).

communication with Kevin Bergersen, 

Arizona Game and Fish Department, 15 Jul 

2002.) 

We speculate that in most of these 

situations the innate attack behavior either 

never was initiated, or was satisfied when 

the puma determined that the prey already 

was moribund, and only needed dragging to 

a protected location to be fed upon. The 

puma simply examines the sleeper, and then 

drags the sleeping bag or mat away from the 

site. When the victim or companions awake 

and begin making noise, the puma is 

frightened and leaves. Perhaps there is an 

element of surprise or ambush by humans; 

perhaps the attack behavior is not initiated 

and therefore does not need to be fulfilled. 

We found, in addition to the 12 mentioned 

above, several oral history accounts (not 

included in our data)of pumas covering 

sleeping people with debris (e.g., Seton 

1929:98-99). 

Are You Safer Riding Horseback? 

Of 9 incidents involving riders on 

horseback, none was seriously injured and 

most were unharmed. In 8 of these cases, 

the puma failed in the initial attack and the 

horse outran it. In the other case, the rider 

killed the puma. If the rider dismounted, 

accidentally or otherwise, the situation 

became more similar to a puma attacking a 

person afoot. 

Risk of Attacks 

Puma attacks on humans are rare by 

almost any measure. Bear attacks are much 

more common. But rarity may be a matter 

of scale, proximity and individuality. Risk 

depends on where people are and what the 

conditions are in that area. Statistics that 

compare the frequency of puma attacks with 

that of some other phenomenon, such as 

lightning strikes, are misleading. Such data 

normally fail to account for risk levels. For 

example, the risk of being attacked by a wild 

puma in a large metropolitan area is close to 

zero, but the large urban areas include most 



of the people who make up the denominator 

in the attack rate. The risk of being attacked 

in a puma’s natural territory is higher, only 

because the puma may be present. The 

same logic applies to bites by domestic 

dogs. Mail carriers are more at risk from 

dog bites than are other people because they 

periodically enter yards that the dogs 

consider to be their territory. In fact, the 

mail carriers’ risk level increases 

dramatically the instant they enter a yard 

containing a dog. 

The situation is similar with puma 

attacks. Most of us have almost no risk, but 

under certain conditions the risk rises 

considerably. Data presented by Mattson et 

al, at the Seventh Mountain Lion Workshop, 

was an effort to predict conceptually some 

of these risk factors. Our presentation, on 

the other hand, is mostly an effort to help 

people manage an attack situation. Data 

presented by Sweanor et al., at the Seventh 

Mountain Lion Workshop, is a strong 

beginning toward collecting data to identify 

risk factors. 

A person in an encounter should always 

judge the puma’s responses and adjust 

defensive measures according to the puma 

response. Each puma has its own 

personality, as the following incident 

illustrates. A United States Forest Service 

employee was in a remote area away from 

roads doing silvicultural analysis. While in 

her sleeping bag in camp she noticed a puma 

at her feet near the edge of her sleeping bag. 

She calmly said, ”excuse me,” whereupon 

the puma moved 4.6 m (15 feet) away and 

sat. It seemed to be a healthy, curious puma. 

If she got upset and tried to make it leave, it 

got aggressive; if she lay there quietly, it 

calmed down, reclined, and licked itself. If 

she yelled at it, it would pin its ears and 

charge. The puma started to circle her, so 

she threw sticks and rocks at it to keep it at 

bay and lit the fire. She called in coordinates 

via radio to her supervisor. The puma


watched all night and finally left the area as 

rescuers arrived. (Personal communication 

from Kathleen Kavalok to K. Etling). 

The incident above illustrates the 

presence of mind needed when confronted 

by a puma. This person did everything just 

right, but the puma responded almost the 

opposite way from what was expected. We 

believe the information we have provided is 

correct in a statistical sense. It is very 

important to remember what to do, but also 

be prepared to adapt to the puma’s behavior. 

With luck and aggressiveness on your part, 

you may avoid an attack and also teach the 

puma that humans may not be food after all. 


We thank S. C. Reed for helping build 

the spreadsheet, verify data, review files, 

and enter new data. J. Schmidt helped with 

the original compilation in 1985-86. R.W. 

Riley, K. J. Stahle, S. E. Gordon, M. A. 

Whittaker, B. R. Campos, M. E. Jackson, E. 

Chen, A. M. White helped with data 

organization and entry. E. L. Blake 

provided some new ideas for analysis. 

Details of incidents were provided by P. 

Swift, California Department of Fish and 

Game, R. Beausoleil, formerly of the New 

Mexico Department of Game and Fish, K. 

Bergersen, Arizona Game and Fish 

Department, T. R. Collom, W. Castillo, and 

D. Whittaker, of the Oregon Department of 

Fish and Wildlife, M. Austin and B. Guiltner 

of the B.C. Ministry of Environment, M. 

Gillett, R. Skiles E. Myers, A. Davis, J. 

Case, K. McKinlay-Jones, of the National 

Park Service, A. Barton, M. Shuey, L. 

Lewis, and S. Galentine. We made 

extensive use of new information in books 

by K. Etling (2001), by J. Deurbrock and D. 

Miller (2001), and by H. P. Danz (1999). 

We thank the many who have provided 

information about their personal 

experiences, either to us or to previous 

authors. P. Beier (1991) provided the first 

scientific analysis of puma attack data, and 


inspired us to continue our work. Finally, 

the very foundation, without which this 

effort may not have ever begun, is C. T. 

Barnes’ (1960) book. His book stimulated, 

and to a large extent, enabled this project 

during its infancy. R. G. Coss, H. G. Shaw, 

L. L. Sweanor, W. F. Laudenslayer, W. E. 

Howard, and R. E. Marsh improved the 

paper with reviews of the draft manuscript, 

but errors remain the author’s responsibility. 


ANONYMOUS. 1917. Untitled comment by 

editor following Marsh 1917. Outdoor 

Life 38:194. 

ANONYMOUS. 1925. An old question. 

Outdoor Life 46:113. 

ANONYMOUS. 1994. Phoenix Gazette and 

Tucson Citizen. 20 July 1994. 

ANONYMOUS. 2002. The New York Times. 

12 November 2002. 

BARNES, CLAUDE T. 1960. "The Cougar or 

Mountain Lion." The Ralton Co. Salt 

Lake City, Utah, USA. 175 pp. 

BEIER, P. 1991. Cougar attacks on humans 

in the United States and Canada. 

Wildlife Society Bulletin 19:403-412. 

COLORADO DIVISION OF WILDLIFE. 2002. 

News Report. 8 January 2002. 

DANZ, HAROLD P. 1999. Cougar! Swallow 

Press/Ohio University Press. Athens, 

Ohio, USA. 310 pp. 

DEURBROCK, JO AND DEAN MILLER. 2001. 

Cat attacks: true stories and hard lessons 

from cougar country. Sasquatch Books. 

Seattle, Washington, USA. 221 pp. 

ETLING, KATHY. 2001. Cougar attacks: 

encounters of the worst kind. The Lyons 

Press/Globe Pequot Press. Guilford, 

Connecticut, USA. 246 pp. 

FITZHUGH, E. LEE AND W.P. GORENZEL. 

1986. Biological status of mountain 

lions in California. pp. 336-346 in 

Proceedings, Twelfth Vertebrate Pest 

Conference. T. P. Salmon (ed.). 

Vertebrate Pest Council of the

Vertebrate Pest Conference. University 

of California, Davis, California, USA. 

FORD, PHIL. 1994. Burney bowhunter 

stalked by fearless mountain lion. 

Fishing and Hunting News. September 

15-29, 1994. 

HUNTER, J. MARVIN. 1922. Pioneer history 

of Bandera County; seventy-five years of 

intrepid history. Hunter’s Printing 

House. Bandera, Texas, USA. 241 pp. 

MARSH, CHARLES. 1917. Children attacked 

by cougar. Outdoor Life 38:193-194. 



PRITCHARD, C. HESKETH. 1902. Through 

the heart of Patagonia. New York, USA. 

346 pp. 

ROOSEVELT, THEODORE. 1914:29. Through 

the Brazilian wilderness. New York, 

USA. 

SETON, E.T. 1929. Lives of game animals. 

Doubleday, Doran & Company, Inc. 

Garden City, New York, USA. Vol I, 

Part I.

104 

A CONCEPTUAL MODEL AND APPRAISAL OF EXISTING RESEARCH RELATED 

TO INTERACTIONS BETWEEN HUMANS AND PUMAS 

DAVID J. MATTSON, USGS Southwest Biological Science Center, Colorado Plateau Field 

Station, P.O. Box 5614, Northern Arizona University, Flagstaff, AZ 86011-5614, USA, 

email: David.Mattson@nau.edu 

JAN V. HART, USGS Southwest Biological Science Center, Colorado Plateau Field Station, 

P.O. Box 5614, Northern Arizona University, Flagstaff, AZ 86011-5614, USA, email: 

Jan.Hart@nau.edu 

PAUL BEIER, School of Forestry, Northern Arizona University, Flagstaff, AZ 86011-5018, 

USA, email: Paul.Beier@nau.edu 

JESSE MILLEN-JOHNSON, Bates College, Lewiston, ME 04240, USA, email: 

jmillenj@bates.edu 

Abstract: Recorded encounters between humans and pumas have been increasing throughout the 

western contiguous U.S., as have puma-caused human injuries and deaths. We developed a 

conceptual model of interactions between humans and pumas to aid the design of a study in the 

Flagstaff uplands of Arizona, USA, and to appraise the scope and strength of existing related 

research. The model represents contact and resulting human injuries as the outcome of 2 

processes: (1) the frequency of encounter between humans and pumas, and (2), given an 

encounter, the probability that it will turn injurious to a human. Conceptually, different suites of 

factors govern these 2 phenomena. The model representing frequency of encounter includes 15 

putative explanatory variables, 7 of which relate directly to pumas. The model representing 

probability of injury also includes 15 explanatory variables, 7 of which pertain directly to pumas. 

The remaining variables in both models relate directly or indirectly to human presence or 

behavior. Of the 44 identified relations among these variables, 6 have been well studied and an 

additional 18 have been subject to some level of systematic analysis. The remaining 20 

relations, including many plausibly critical ones, are currently informed only by speculation, 

anecdote, or deduction. Much research yet needs to be done before the level and nature of 

contact between humans and pumas can be adequately explained and predicted. Moreover, 

much of this additional research needs to address human behavior and factors related to 

distributions and numbers of humans. Of the uninvestigated factors with plausibly major effects, 

habituation of cougars promises to be the most difficult to study. Otherwise, numbers and 

distributions of human facilities (including roads and trails), puma population sizes, human 

behavior, and human knowledge of pumas are potentially important explanatory factors 

amenable to inquiry. We argue that all of these putative effects should be considered before 

reaching conclusions about the “causes” of human-puma encounters and puma-caused human 

injuries, whether for a region or a given study area. 


RELATIONSHIPS BETWEEN LAND TENURE SYSTEM, MOUNTAIN LION 

PROTECTION STATUS, AND LIVESTOCK DEPREDATION RATE 

MARCELO MAZZOLLI, Projeto Puma - R. Liberato Carioni 24, Lagoa, 88062-005, 

Florianópolis – SC, Brazil, email: marcelo_puma@yahoo.com 

Abstract: Mountain lion depredation impact on managed livestock in ranching-dominated 

landscapes was compared with depredation in forestry-dominated landscapes. In forestrydominated 

landscapes, twenty-one depredation incidents were recorded at three ranches, 

resulting in the loss of 58 sheep and goat. An additional 11 head were killed during an unknown 

number of attacks. These losses amounted to 14 to 60 percent of total stock per year (in number 

of animals). Confining or corralling flocks during the night at first provoked a reduction of 

livestock depredation, but subsequently depredation begun to occur during daylight. In the 

ranching-dominated landscape, on the other hand, depredation losses were not reported on flocks 

corralled during the night, but reached 85% of the total stock when free-ranging. I hypothesize, 

based on field data and on available information in literature, that mountain lion depredation in 

forestry areas during the day and higher depredation on corralled livestock during the night may 

result from lower hunting pressure on mountain lions than in ranching areas. Understanding 

mountain lion predation behavior may help to modify livestock husbandry, allowing wildlife 

managers and ranchers to minimize depredation without direct persecution of mountain lions. 


105

106 

MOUNTAIN LION MOVEMENTS AND PERSISTENCE IN A FRAGMENTED, URBAN 

LANDSCAPE IN SOUTHERN CALIFORNIA 

SETH P.D. RILEY, Santa Monica Mountain National Recreation Area, 401 W. Hillcrest Dr., 

Thousand Oaks, CA 91360, USA, email: seth_riley@nps.gov 

RAYMOND M. SAUVAJOT, Santa Monica Mountain National Recreation Area, 401 W. 

Hillcrest Dr., Thousand Oaks, CA 91360, USA 

ERIC C. YORK, Santa Monica Mountain National Recreation Area, 401 W. Hillcrest Dr., 

Thousand Oaks, CA 91360, USA, email: eric_york@nps.gov 

Abstract: As natural habitat is increasingly eliminated and fragmented by human land uses the 

long-term prospects for conservation of carnivore populations become correspondingly worse. 

This is especially true for larger carnivores such as mountain lions, which require significant 

amounts of both space and prey. In rapidly urbanizing southern California, conservation of 

carnivores in general, and of mountain lions in particular, is particularly challenging. In the 

Santa Monica Mountains and surrounding areas, we have begun a project using GPS collars to 

determine mountain lion movement and space use in a fragmented landscape. Our goal is to 

determine whether lions are successfully traversing freeways and other human-made barriers 

between large areas of natural habitat. Ultimately, we hope to determine whether enough natural 

habitat can be preserved, and enough connectivity maintained between core habitat areas, to 

maintain lion populations in such a landscape. We have collared lions already in the Santa 

Monica Mountains, and determined that one large male is using the entire mountain range (home 

range of 394 km 2 ), from a major freeway to the east to a developed agricultural valley to the 

west, and from the Pacific Ocean to the south to a major freeway to the north. Given the small 

number of lions likely persisting in the Santa Monica Mountains, connectivity is as important, if 

not more important, than we anticipated. We continue to collar other lions in the study region to 

evaluate whether any exchange occurs across barriers created by freeways and urban 

development. While both the male and the female in the Mountains have approached the 

freeway to the north, neither one has crossed it in the 9-12 months that we have been following 

them. We are also investigating kill sites to determine kill rates, species of kills, and whether 

lions are preying on any domestic animals. So far in the Santa Monica Mountains our data 

indicate that lions are killing 3-4 deer/month of all different age/sex classes, and an occasional 

coyote or raccoon. The collared animals are almost never seen by anyone, including the 

researchers tracking them, even though they cross numerous roads and trails and sometimes 

venture close to residential areas. 


PUMA RESPONSES TO CLOSE ENCOUNTERS WITH RESEARCHERS 

LINDA L. SWEANOR, Wildlife Research Biologist, Wildlife Health Center, University of 

California at Davis, Southern California Puma Project Field Station, P.O. Box 1114, 

Julian, CA 92036-1114, USA, email: lsweanor@mindspring.com 

KENNETH A. LOGAN, Wildlife Research Biologist, Wildlife Health Center, University of 

California at Davis, Southern California Puma Project Field Station, P.O. Box 1114, 

Julian, CA 92036-1114, USA 

MAURICE G. HORNOCKER, Senior Scientist, Wildlife Conservation Society, Box 929, 

Bellevue, ID 83313, USA 

Abstract: Recent books and articles have provided information on relatively rare, but violent 

attacks where people were injured or killed by pumas. However, there is a paucity of data on the 

type and variation in behavior wild pumas exhibit when approached by humans. During a 10year 

puma study in New Mexico, we approached pumas and visually observed their behavior on 

262 occasions. The study area was remote and was closed to most human activity; consequently 

the pumas living there had rare opportunities for contact with people. We categorized the 

approach based on the status of the puma, the number of people involved, the distance between 

the puma(s) and people, and the puma’s subsequent response to the approach. Pumas we 

approached included adult females with nursing (

108 

PRELIMINARY RESULTS OF FLORIDA PANTHER GENETIC ANALYSES 

WARREN E. JOHNSON, National Cancer Institute, Frederick, MD 21702-1201, USA, email: 

johnsonw@ncifcrf.gov 

DARRELL LAND, Florida Fish and Wildlife Conservation Commission, 566 Commercial 

Blvd., Naples, FL 34104, USA, email: darrell.land@fwc.state.fl.us 

JAN MORTENSON, National Cancer Institute, Frederick, MD 21702-1201, USA, email: 

martenoj@ncifcrf.gov 

MELODY ROELKE-PARKER, National Cancer Institute, Frederick, MD 21702-1201, USA, 

email: roelkem@ncifcrf.gov 

STEPHEN J. O’BRIEN, National Cancer Institute, Frederick, MD 21702-1201, USA, email: 

obriens@ncifcrf.gov 

Abstract: Previous genetic analyses showed that Florida panthers (Puma concolor coryi) had the 

lowest genetic diversity among all North American puma and subsequent modeling suggested 

that further declines could increase the probability of extinction. Currently, there are fewer than 

100 panthers in south Florida. Although on-going habitat conservation strategies may provide 

long-term stability for today’s population extents, these same strategies are unlikely to allow the 

population to grow to 500 or more individuals whereby genetic viability is more assured. As a 

result, a plan for Florida panther genetic restoration was created in 1994 and implementation 

began in the spring of 1995 with the release of 8 female Texas puma into areas occupied by 

panthers. Our objectives were to monitor the effectiveness of genetic restoration by developing 

an array of molecular genetic markers that characterized the status of current and past 

populations, to construct a pedigree among Florida panthers to follow inheritance patterns, to 

infer degrees of relatedness among individuals, and to help predict the future viability of the 

population. We have completed genotyping over 175 samples from Florida panthers at 23 

microsatellite loci and these included individuals from canonical Florida panthers, the 

Everglades subpopulation (Piper stock), released Texas puma, crosses among all stocks, and 

captive animals of unknown ancestry from the early 1970’s to the present. Genetic restoration 

has increased hetereozygocity within the population, but we have documented the loss of some 

panther matrilines. Certain morphological traits such as cryptorchidism, kinked tails, cowlicks, 

and atrial septal defects observed in canonical panthers are not present in the Texas puma 

descendants. We have identified several subgroups within our population and these subgroups 

seem to be partially the product of philopatric tendencies among dispersing female offspring. 

Male panthers may be physically and behaviorally capable of siring offspring earlier than 

suggested by radiotelemetry work and resident and resident males are not siring all litters with 

females within the respective males’ home ranges. Intraspecific aggression, a common mortality 

agent for young male panthers, may not be removing panthers prior to producing offspring. 

Future monitoring should ensure sampling across all panther subgroups in order to adequately 

estimate total population genetic characteristics. 


GENETIC STRUCTURE OF COUGAR POPULATIONS ACROSS THE WYOMING 

BASIN: METAPOPULATION OR MEGAPOPULATION 

CHUCK R. ANDERSON, JR., Zoology and Physiology Department, University of Wyoming, 

Box 3166, University Station, Laramie, WY 82071, USA, email: cander@uwyo.edu 

FRED G. LINDZEY, Wyoming Cooperative Fish and Wildlife Research Unit, Box 3166, 

University Station, Laramie, WY 82071, USA, email: flindzey@uwyo.edu 

DAVE B. McDONALD, Zoology and Physiology Department, University of Wyoming, 

Bioscience Room 413, University Station, Laramie, WY 82071, USA, email: 

dbmcd@uwyo.edu 

Abstract: Using microsatellite DNA analyses at 9 loci, we examined genetic structure of 5 

geographically distinct cougar (Puma concolor) populations separated by the Wyoming Basin 

and a distant cougar population from southwest Colorado. Observed heterozygosity was similar 

among populations (Hobs = 0.49-0.59) and intermediate to that of other large carnivores. 

Estimates of genetic structure (FST = 0.029, RST = 0.028) and number of migrants per generation 

(Nem) suggested high gene flow across the central Rocky Mountains. Estimates of the number of 

migrants per generation were lowest between the southwest Colorado cougar population and 

cougar populations north of the Wyoming Basin (northwest WY, north-central WY, and the 

Black Hills, SD; Nem = 2.9-3.0) and highest among cougar populations from adjacent mountain 

ranges (Nem = 10.2-30.2), suggesting an effect of both isolation by distance and of habitat 

matrix. We applied a model-based clustering method to infer population structure from 

individual genotypes and noted that both males and females from throughout the region were 

best described as a single panmictic population. Estimates of relatedness (rxy) did not differ (P > 

0.05) between males and females. Estimated relative effective population size did not differ 

significantly among populations (P > 0.05), but the higher estimates were from contiguous 

mountain ranges (i.e., northwest WY, southwest WY, and southwest CO) and lower estimates 

were from less contiguous terminal mountain ranges (i.e., north-central WY and Snowy Range 

WY). Based on measures of gene flow we examined, extinction risk in the near future appears 

extremely low, even for the relatively isolated Black Hills cougar population. Cougars in the 

central Rocky Mountains appear to constitute a large panmictic population rather than a 

metapopulation. Estimated effective population size for cougars in the central Rocky Mountains 

ranged from 1,797 to 4,532 depending on analysis method and the mutation model assumed. 


109

110 

ECOLOGICAL SIGNIFICANCE AND EVOLUTION OF A COMMON COUGAR 

RETROVIRUS 

ROMAN BIEK, Fish and Wildlife Biology Program, University of Montana, Missoula, MT 

59812, USA, email: rbiek@selway.umt.edu 

MARY POSS, Fish and Wildlife Biology Program and Division of Biological Sciences, 

University of Montana, Missoula, MT 59812, USA, email: mposs@selway.umt.edu 

Abstract: As for most wildlife species, little is known about the organisms that infect cougars in 

the wild. In an ongoing project, we are studying a retrovirus related to domestic cat-FIV that is 

naturally found in North American cougars with the aim of assessing the virus’ possible 

demographic consequences on the cougar host as well as its epidemiology and short-term 

evolution. Tests for possible effects on survival and reproduction as well as secondary exposure 

to other pathogens in infected individuals are conducted based on a large data set compiled from 

several intensively studied cougar populations. In addition, DNA sequences of virus obtained 

from infected individuals are used to determine the genetic population structure of cougar-FIV in 

the Rocky Mountain region. We determined that that this virus is changing its genetic 

composition within a matter of decades. Because restrictions of cougar movement will be 

reflected in the distribution of closely related viruses, distributional data for the virus are thus 

likely to contain information about current patterns of connectivity among cougar populations. A 

preliminary analysis of these data indicates that spread of the virus occurs mainly locally but also 

showed evidence for recent transmission events over distances > 300 km. These results show that 

studying the ecology of cougar-FIV can provide important insights into the ecology of the cougar 

host even beyond immediate disease impacts. 


CHARACTERISTICS OF MOUNTAIN LION BED, CACHE AND KILL SITES IN 

NORTHEASTERN OREGON 

JAMES J. AKENSON, Taylor Ranch Field Station, HC 83 Box 8070, Cascade ID 83611, USA, 

email: tayranch@directpc.com 

M. CATHY NOWAK, Cat Tracks Wildlife Consulting, P.O. Box 195, Union, OR 97883, USA, 

email: mcnowak@eoni.com 

MARK G. HENJUM, Oregon Department of Fish and Wildlife, 107 20 th Street, La Grande, OR 

97850, USA 

GARY W. WITMER, USDA APHIS National Wildlife Research Center, 4101 La Porte Avenue, 

Fort Collins, CO 80521, USA 

Abstract: We described mountain lion (Puma concolor) habitat characteristics during two studies in the same area of 

northeastern Oregon during the 1990s. In the first study (1992-1994) we evaluated micro-habitat features associated 

with 61 diurnal bed sites that were not associated with kills. We used similar techniques in the second study (1996- 

1998) to evaluate habitat features at 79 cache sites near lion-killed prey. A dog was used to find 93% of the diurnal 

bed sites. Radio telemetry triangulation was used in the second study. Characteristics of diurnal bed sites and cache 

sites were compared with random habitat plots. Rock structure and downed logs were identified as important habitat 

components at diurnal bed sites. Canopy cover at cache sites was significantly higher than at random sites. Cache 

sites also were associated with rock structure, but not to the same degree as diurnal bed sites. In both studies 

mountain lions used sites in close proximity to habitat edges more frequently than expected based on random plots. 

Understanding the similarities and differences of habitat use at diurnal bed, cache and kill sites sheds light on the 

ecological adaptation of mountain lions to the multiple environmental influences and disturbances of managed 

forests. 

111 


Key words: Puma concolor, microhabitat use, diurnal bed site, cache site, kill site, habitat edge, forest management 

Mountain lion recovery has been one of 

the great wildlife conservation success 

stories of the 20 th century. As we move into 

the 21 st century, the challenges for mountain 

lion conservation are less related to species 

persecution, and more related to concerns 

with habitat fragmentation and issues of 

human-lion coexistence on the expanding 

fringe of urbanization. The interface 

between human resource development and 

mountain lion habitat use has persisted for 

centuries in North America. Historically, 

mountain lions have occupied most habitats 

occurring on this continent. Mountain lions 

have typically been associated with the 

rugged, rocky, forested terrain of the Rocky 

Mountains in the western United States; 

however, this species is so adaptable it can 

thrive in deserts, swamps, tropical jungles, 

and sub-alpine forests (Hornocker 1976). 

The lion has come into conflict with humans 

on several fronts. In the past, the majority 

of interactions between humans and 

mountain lions were associated with 

settlement and agricultural practices (Young 

1946). With increasing human population 

and urban sprawl, the zone of conflict has 

shifted to the urban-wildland interface 

(Beier 1995). 

Habitat fragmentation can take a more 

subtle form than the direct effect imparted 

by urbanization. Across much of the 

mountain lion’s range, logging has occurred 

at various intensities. Studies in Utah and 

Arizona, found that mountain lions either 

avoided active timber sale areas (Van Dyke

112 MOUNTAIN LION BED, CACHE AND KILL SITES · Akenson et al. 

et al. 1986) or adjusted their activity pattern 

from the norm (Ackerman 1982), to 

maximize night-time concealment from 

human contact. Timber sale size, relative to 

a resident mountain lion’s home range, was 

a big factor on the degree of disturbance and 

influence on a lion’s willingness to maintain 

its home range (Van Dyke et al. 1986). 

Small-area logging operations were less of a 

negative factor for resident adults. Van 

Dyke et al. (1986) also concluded that 

dispersing young animals were more 

adversely affected by logging and road 

system development than were established 

adults. By comparison, Gagliuso (1991) did 

not find avoidance by radio-collared lions to 

either recent logging or high road densities 

in his southwestern Oregon study area. 

Differences in his findings from Van Dyke 

et al. were related to under-story density and 

rapid recovery of brush in newly logged 

areas. The southwest Oregon study area had 

more than twice the precipitation of the 

Arizona and Utah studies. 

We compare the results of our studies 

within the same northeast Oregon study area 

and discuss similarities and differences with 

studies in Utah, Arizona and southwest 

Oregon (Van Dyke et al. 1986, Gagliuso 

1991). Our studies in northeast Oregon 

were conducted in a climatological, 

geographical, and anthropogenic situation 

somewhere in-between those areas described 

by Van Dyke et al., and Gagliuso. The 

objectives of this paper are to: 1) connect 2 

habitat investigations to gain a more 

complete understanding of microhabitat use 

relative to mountain lion life history, and 2) 

compare mountain lion microhabitat use in 

northeast Oregon with similar work in other 

regions of the western United States. 

STUDY AREA 

Both of these studies were conducted in 

the Catherine Creek Wildlife Management 

Unit in northeast Oregon. The Catherine 

Creek study area is approximately 845 km 2 


in size. Elevations range from 940 to 2,450 

m. This area is flanked on the west by range 

and agricultural lands of the Grande Ronde 

Valley and on the east by the Wallowa 

Mountains within the Eagle Cap Wilderness 

Area. Most of the area (60%) is on the 

Wallowa Whitman National Forest, with the 

remaining being divided between Boise 

Cascade Corporation lands and other private 

ownership. Vegetation varies from 

subalpine coniferous forest to mixed conifer 

forest to rangeland and cropland. Road 

density varies from medium-high density 

(1.4 km/km 2 ) to small road closure areas. 

Approximately 20% of the work from these 

studies was conducted within a Boise 

Cascade road closure area that had received 

various levels of logging activity. The 

majority of this area is mid-elevation 

coniferous forest with various forms of rock 

structure including rimrocks and outcrops. 

METHODS 

We compared the primary findings of 

habitat characteristics at diurnal bed sites in 

Akenson et al. (1996) and at kill and cache 

sites in Nowak (1999). The 2 studies were 

compared qualitatively and the similarities 

and differences were described and 

discussed in an ecological context. The 

methods utilized in the 2 studies are briefly 

described below. 

Akenson et al. (1996) used various 

methods of locating and identifying 

mountain lion diurnal bed sites including 

snow tracking, radio telemetry, and a trained 

lion hound that located scent at bed sites. 

These methods were modified from 

Anderson (1990) for locating bobcat loafing 

sites in Colorado. A bed site was confirmed 

through visible evidence of either soil or 

litter disturbance or tracks, and by alert 

reactions of a reliable dog. Beds were 

typically visible as a depression in snow or 

duff, or flattened grass. Once a bed site was 

identified, the surrounding area was 

searched for prey remains to determine

whether the bed was associated with a kill. 

The actual bed site became the center of a 

50-meter radius plot for collection of data to 

determine the physiographic and vegetative 

composition of the site. Habitat descriptions 

were aided by the handbook “Plant 

Associations of the Wallowa - Snake 

Province” (Johnson and Simon 1987). 

Akenson et al. (1996) evaluated 6 

primary habitat features at each plot site 

including rock structure, forest structure, 

canopy cover, shrub cover, plot visibility 

and overall security from human 

disturbance. This study emphasized the 

structural influence of vegetation and 

topography on a mountain lion’s security 

from detection. Other environmental 

influences such as distance to roads and 

abrupt habitat edges were also recorded. 

The distance to road measurement was 

recorded from the plot center to the nearest 

drivable road. A habitat edge typically 

marked a forest break or the beginning of a 

rock wall or large rock outcrop. For 

comparison, habitat data were also collected 

at randomly selected sites distributed 

throughout the study area. Random sites 

corresponded to the same square-mile 

section corner in 30 sections drawn from a 

pool of 185 possibilities, which all occurred 

in the known home ranges of the 5 subject 

mountain lions. All mountain lion age and 

sex classes were included. Habitat plots 

were categorized as summer (April 15 to 

September 1), winter (December 15 to 

March 15) or random, and data were 

summarized and compared using chisquared 

tests for differences between the 3 

plot types. Values were considered 

significant at α = 0.05. 

Nowak (1999) applied the term “cache 

site” to the location where a mountain lion 

kill was first found, whether or not the lion 

had moved it after making the kill. The 

exception to this was if the kill had 

obviously been moved from the original 

MOUNTAIN LION BED, CACHE AND KILL SITES · Akenson et al. 113 


cache site for subsequent feeding and the 

original cache site could be identified. The 

term “kill site” referred to the location 

where the mountain lion actually killed its 

prey. The distinction between cache and kill 

site involved a combination of telemetry 

triangulation when the lion was present, and 

then an investigation of the area after the 

lion moved a safe distance away. As with 

other studies on lions, the majority of 

information was obtained from locating 

radio instrumented animals on the ground 

(Anderson et al. 1992). Once the cache or 

kill site was determined, then this site 

became the center of a 25-meter radius plot 

for collection of physiographic and 

vegetative data. 

Work closely followed Akenson et al. 

(1996) to facilitate comparisons between the 

2 studies. Data were collected for 25 habitat 

variables to evaluate rock structure, forest 

structure, canopy cover, plot visibility and 

proximity to potential disturbance. This 

study likewise emphasized the influences of 

vegetation and topography on mountain lion 

security but also on the security of kills, 

which may be left unattended for long 

periods of time. In this study, distance was 

recorded to both the nearest open, drivable 

road and to the nearest road of any kind, 

open or closed. As with Akenson et al., a 

habitat edge was typically a relatively abrupt 

change in stand composition and/or structure 

or topography. For comparison, habitat data 

were also collected at randomly selected 

sites distributed throughout the study area 

but within the subject lions’ home ranges. 

UTMs for random plots were generated by a 

computer random number generator 

(Microsoft Excel) using known study animal 

home ranges as limits to the generated 

coordinates. Habitat plots were categorized 

as cache, kill or random, and data were 

summarized and compared using forward, 

stepwise, logistic regression for differences 

between the 3 plot types. Values were


considered significant at α = 0.05. Only 

adult female mountain lions, with and 

without young, were included. 

RESULTS 

Akenson et al. (1996) recorded habitat 

characteristics at 61 diurnal bed sites, 32 

during winter and 29 during summer. Most 

(87%) of these sites were not associated 

with kills. They collected the same habitat 

data at 30 randomly selected plots. Nowak 

(1999) collected habitat data at 79 cache 

sites, 19 kill sites and 101 randomly selected 

sites. 

Akenson et al. (1996) found significant 


differences between diurnal bed sites and 

randomly selected sites in presence of rock 

structure, number of down logs in the plot, 

distance to habitat edge, sight distance (the 

median distance at which a person could be 

seen from plot center at about lion height), 

understory density and management status 

(Table 1). Nowak (1999) found significant 

differences between cache sites and 

randomly selected sites in canopy cover, 

understory density, elevation, and 

management status. Significant differences 

between kill and random sites were in 

elevation, management status and plot 

visibility (the mean distance at which a 

Table 1. Habitat characteristics at mountain lion diurnal bed sites, summer and winter, at cache 

sites, and at randomly selected sites associated with each study (Akenson et al. 1996, Nowak 1999). 

Asterisks (*) indicate features significantly different (p

person could be seen from plot center at 

about lion height). Kill and cache sites 

differed only in canopy cover (Table 1). 

Large rock structure (forested rimrock) 

and down logs were present in significantly 

more diurnal bed site plots than expected but 

that was not the case for cache sites, 

although cache sites were slightly more 

likely to contain rock ledges than were the 

random sites in that study. Canopy cover 

was significantly greater in cache sites than 

in either kill or random sites but was not 

different between bed sites and random 

sites. Understory density was lower in 

cache sites but higher in summer diurnal bed 

sites. Akenson et al. (1996) found greater 

use of the old logged management type for 

diurnal beds in winter; Nowak (1999) found 

cache sites in old logged with similar 

frequency to random plots (Table 2). A 

relatively high percentage of cache sites 

were located in shelterwood but diurnal beds 

were in that management type with similar 

frequency to random plots. Cache sites were 

in the rangeland management type with less 

frequency than the random sites but bed 

sites were located in rangeland with about 

the same frequency as random sites. 

Neither study documented significant 

differences in the distance to the nearest 

open road although both winter bed sites and 

cache sites tended to be farther from open 

roads than random sites. In Akenson et al. 



(1996), summer diurnal beds were 

significantly closer to a habitat edge than 

were random sites. Although not 

statistically significant, Akenson et al. 

(1996) and Nowak (1999) found that winter 

diurnal beds and cache sites both tended to 

be closer to a habitat edge than the random 

sites. Both studies documented significantly 

lower plot visibility/sight distance in sample 

plots compared with random sites. Both 

studies also showed seasonal variation, in 

elevation with both bed sites and caches at 

lower elevation in winter than in summer. 

When 4 seasons were considered, Nowak 

found cache sites were at higher elevation in 

fall than in summer, spring, or winter. 

DISCUSSION 

Several authors have addressed the 

question of mountain lion habitat use, 

conducted studies in some diverse 

environments, and concluded that a primary 

factor in habitat selection for this carnivore 

was the presence of vegetation and terrain 

cover to enhance the stalking of prey, 

usually deer or elk. Hornocker (1970) felt 

that lions in his Idaho study area selected 

habitat on the basis of prey density and 

terrain features that were advantages for 

hunting. Logan and Irwin (1985) also noted 

a high occurrence of lion caches within 

canyon vegetation, draws, and on steep 

ridges demonstrating the importance of both 

Table 2. Management status at mountain lion diurnal bed sites, summer and winter, at cache sites, 

and at randomly selected sites associated with each study (Akenson et al. 1996, Nowak 1999). 

Asterisks (*) indicate features significantly different (p


vegetative and terrain cover. Seidensticker 

et al. (1973) concluded that a “vegetation – 

topography/prey numbers – vulnerability 

complex” determined both lion home range 

size and population density. We agree that 

the need for cover while bedding, hunting, 

or guarding a cache site is ecologically 

important. Our findings indicate that forest 

management strategies contribute to both 

prey abundance and enhanced stalking cover 

for mountain lions (Table 2). 

Van Dyke et al. (1986) concluded that 

resident lions avoided portions of their home 

ranges with active logging activity, and 

found that transient lions were the primary 

users of areas with active timber harvest, or 

even newly logged areas. By contrast, 

Gagliuso (1991) found in southwestern 

Oregon that lions did not avoid timber 

harvest sites but rather were closer to these 

activities than expected at random. We 

observed a similar attraction to new logging, 

which we believed was related to the 

abundant “candy food” made newly 

available to deer and elk by logging that 

brought branches laden with lichen and 

mosses down to ground level. Once this 

resource was exhausted, deer and elk quit 

using these sites, as did hunting lions. We 

concluded from track evidence made in 

snow during winter, or dust during summer, 

that lions were using newly logged areas at 

night. Nocturnal movement patterns, in 

association with sub-optimal habitat cover, 

was also documented by Beier (1995) in 

California and Van Dyke et al. (1986) in 

Utah where they documented mountain lions 

using the most undisturbed habitats in their 

home ranges for diurnal localization. Our 

findings concur with these authors. On a 

micro-habitat scale, our findings also show 

the importance of specific features, such as 

forested rimrock and downed logs for 

diurnal bed sites, understory density for 

hunting and stalking cover, and canopy 

cover for kill cache sites. 


The documentation of micro-habitat use 

is essential in understanding mountain lion 

daily adaptation to multiple environmental 

influences and disturbances. The use of 

specific habitat types by lions is largely 

dependent on the activity of the individual. 

A cougar that is bedding for the day selects 

a location that offers both concealment and 

nearby escape terrain, as indicated in our 

study by a strong selection for forested 

rimrock structure with a component of 

downed logs. Whereas a lion that is hunting 

is going to use areas preferred by prey 

species that also afford stalking 

concealment, usually in the form of understory 

vegetation or other close to the ground 

structure. Then, once the kill has been 

made, there is typically an effort made by 

the lion to cache the kill under a tree or 

brush, presumably to reduce detection by 

avian scavengers. 

MANAGEMENT IMPLICATIONS 

Our findings on mountain lion habitat 

use have implications to both wildlife and 

habitat managers. There are many complex 

variables influencing mountain lion habitat 

use in different regions and levels of human 

influence. Several factors influence the way 

in which lions use their environment, or 

conduct “land tenure” as described by John 

Seidensticker (1973). Obtaining food, 

establishing and defending territories, 

breeding, reproducing, and raising kittens to 

dispersal age all have a bearing on how 

mountain lions use a given landscape. In 

comparing findings from this study with 

other studies, it appears that factors vary 

from region to region. However, habitat use 

seems to be driven by three ecological 

needs: security, cover, and food. 

The mountain lions that we studied have 

co-existed with timber harvest for several 

lion generations. The literature suggests that 

lions will still use habitats that have been 

logged as long as the harvest areas are

Gagliuso 1991). Leaving strips of trees for 

buffers, in conjunction with small harvest 

units, creates an extensive habitat edge 

effect beneficial to mountain lions. Other 

important features are vegetative cover 

around rock structure for bedding security, 

downed logs, and ample understory density 

to allow for successful stalking. All of the 

diurnal bed-sites occurring in rimrock had 

either brush or trees at the bed. We did not 

document bed-use in newly logged areas or 

in rock structure without some form of 

vegetative cover. A timber management 

practice that leaves a forested buffer around 

rock structure is advantageous for mountain 

lion security. The size of the buffer would 

vary with vegetation type and density, but 

generally a 50-meter buffer would afford 

concealment for lions in our study area. We 

did not find a significant aversion to roads in 

the Catherine Creek study area, but our 

methods may not have effectively addressed 

this issue since most of our data was 

gathered in or near a Boise Cascade 

Corporation road closure area. The two 

primary land managers, the US Forest 

Service and Boise Cascade Corporation, 

have implemented travel management plans 

that vastly reduce human disturbance 

through established road closure areas. In 

general, our findings are more similar to 

results produced in southwest Oregon by 

Gagliuso (1991) than those described by 

Van Dyke et al. (1986) in Arizona and Utah. 

We feel these differences are due to 

mountain lions in Oregon having long-term 

exposure to logging, and the habitat having a 

quicker capability for regrowth with higher 

amounts of precipitation in two areas of 

Oregon than the more arid Southwest. 

SUMMARY 

In conclusion, we have added more 

information to the pool of knowledge 

supporting the concept of mountain lions as 

an adaptable, yet vulnerable species. Logan 



and Sweanor (2001) emphasize the 

importance of gaining a better understanding 

of mountain lion habitat use through 

identifying critical habitats, landscape 

linkages, and by assessing how human 

development, resource extraction, and 

habitat modification can degrade or enhance 

these habitats. We have demonstrated the 

importance of small-scale physiographic 

features within the larger scale habitat 

complex. Scientific management of 

mountain lions depends on both wildlife 

managers and land managers understanding 

this species’ requirements of security, cover, 

and food, and how obtaining these 

ecological needs varies between regions and 

physiographic and climatological situations 

and conditions. 


We are indebted to several agencies and 

individuals who made our studies 

successful. The study by Akenson et al. was 

for the Oregon Department of Fish and 

Wildlife. Funding was mostly from a grant 

from the Federal Aid in Fish and Wildlife 

Restoration Act. The generous contributions 

of time, knowledge, and hound services by 

Ted Craddock, Gale Culver, Loren Brown 

and field assistant Paul Alexander were 

invaluable. Financial support for the study 

by Nowak came from Washington State 

University, the National Wildlife Research 

Center of the U.S.D.A. Animal and Plant 

Health Inspection Service, and the Oregon 

Department of Fish and Wildlife. Nowak is 

greatly indebted for assistance from her field 

crew of: Craig Whitman, Gail Collins, Brett 

Lyndaker, Jeff Olmstead, Tracy Taylor, and 

volunteers: Renan Bagley, Doug Wolf, 

Mark Berrest, Kate Richardson, Keith 

Wehner, Eric Macy, and Mark Squire. The 

authors wish to thank Kerry Murphy and 

Mark Penninger for their constructive 

comments on an earlier version of the 

manuscript.



ACKERMAN, B.B. 1982. Cougar predation 

and ecological energetics in southern 

Utah. Thesis, Utah State University, 

Logan, Utah, USA. 

ANDERSON, A.E., D.C. BOWDEN, AND D.M. 

KATNER. 1992. The Puma on the 

Uncompahgre Plateau. Colorado 

Division of Wildlife. Tech. Pub. No. 40. 

ANDERSON, E.M. 1990. Bobcat diurnal 

loafing sites in southeastern Colorado. 

Journal of Wildlife Management. 

54:600-602. 

AKENSON, J.J., M.G. HENJUM, AND T.J. 

CRADDOCK. 1996. Diurnal bedding 

habitat of mountain lions in northeast 

Oregon. Abstract in Fifth Mountain 

Lion Workshop, 27 February-1 March, 

1996, San Diego, California. 

BEIER, P. 1995. Dispersal of juvenile 

cougars in fragmented habitat. Journal 

of Wildlife Management. 59:228-237. 

GAGLIUSO, R.A. 1991. Habitat alteration 

and human disturbance: their impact on 

cougar habitat utilization in southwest 

Oregon. Thesis, Oregon State 

University, Corvallis, Oregon, USA. 

HORNOCKER, M.G. 1970. An analysis of 

mountain lion predation upon mule deer 

and elk in the Idaho Primitive Area. 

Wildlife Monograph No. 21: 1-39. 

_____. 1976. Cougars up close. National 

Wildlife. 14(6):42-47. 

JOHNSON, C.G. AND S.A. SIMON.1987. Plant 

Associations of the Wallowa-Snake 


Province. U.S. Forest Service handbook 

No. R-6 ECOL-TP-225B-86. 

LOGAN, K.A. AND L.L. IRWIN. 1985. 

Mountain lion habitats in the Bighorn 

Mountains, Wyoming. Wildlife Society 

Bulletin 13:257-262. 

LOGAN, K.A. AND L.L. SWEANOR. 2001. 

Desert Puma, evolutionary ecology and 

conservation of an enduring carnivore. 

Island Press. Washington, D.C., USA. 

NOWAK, M.C. 1999. Predation rates and 

foraging ecology of adult female 

mountain lions in northeastern Oregon. 

Thesis. Washington State University, 

Pullman, Washington, USA. 

SEIDENSTICKER, J.C. IV, M.G. HORNOCKER, 

W.V. WILES AND J.P. MESSICK. 1973. 

Mountain lion social organization in the 

Idaho Primitive Area. Wildlife 

Monograph 35:1-60. 

SEIDENSTICKER, J.C. IV. 1973. Mountain 

lion social organization in the Idaho 

Primitive Area. Dissertation, University 

of Idaho, Moscow, Idaho, USA. 

VAN DYKE, F.G., R.H. BROCK, H.G. SHAW, 

B.B. ACKERMAN, T.P. HEMKER AND 

F.G. LINDZEY. 1986. Reactions of 

mountain lions to logging and human 

activity. Journal of Wildlife 

Management, 50:95-102. 

YOUNG, S.P. 1946. History, life habits, 

economic status, and control, Part 1. 

Pages 1-173 in S.P. Young and E.A. 

Goldman, eds. The puma, mysterious 

American cat. The American Wildlife 

Institute, Washington, D.C. USA.

IMPACT OF EDGE HABITAT ON HOME RANGE SIZE IN PUMAS 

JOHN W. LAUNDRÉ, Instituto de Ecología, A.C. Apartado Postal 632, 34100 Durango, Dgo., 

México, email: launjohn@prodigy.net.mx 

LUCINA HERNÁNDEZ, Instituto de Ecología, A.C. Apartado Postal 632, 34100 Durango, 

Dgo., México, email: lucina@sequia.edu.mx 

Abstract: In the previous workshop in San Antonio, researchers from Wyoming reported that 

pumas from two areas with different amounts of fragmentation still had home range areas that 

contained equal amounts of periphery (= edge). In the same workshop, we reported that edge 

habitat was critical for successful hunting of deer by pumas. These two results indicate that the 

amount of edge habitat in an area may be an important factor in determining home range size of 

pumas. We tested this hypothesis with data we have on home ranges of pumas in southern 

Idaho/northwestern Utah. The study area is highly fragmented into forest patches and sagebrush 

open areas. We tested three predictions: 1) the amount of edge habitat in the home ranges of 

pumas would be similar, regardless of the size of the home range, 2) the percent of edge would 

be negatively related to home range size, and 3) there would be more edge habitat within home 

range boundaries than in general areas of similar size. We tested these predictions by overlaying 

telemetry locations on habitat maps of the area, determining the home range boundaries with the 

minimum convex polygon method and then estimating the amount of forest edge (km 2 ) that 

occurred in each home range. The analysis was conducted with standard GIS software and we 

had 20 pumas where the home range was adequately determined (> 30 relocations). Home 

range size varied from 38 to 120 km 2 . However, 14 (70%) of the home ranges were between 38 

to 105 km 2 . The amount of edge habitat within all the home ranges varied from 13 to 35 km 2 . 

Within the 14 smaller home ranges, the amount of edge varied from 13 to 20 km 2 . The percent 

of edge within home ranges was negatively correlated with home range size. The amount of 

edge within the home range boundaries was significantly greater (F = 15.05, P < 0.001) than 

general areas of similar size. We concluded that the amount of edge within an area was 

influencing the size of home ranges. We proposed that pumas needed a certain minimum 

amount of edge (hunting habitat) to successfully hunt their prey and that the amount of 

“catchable” prey was more important than just general prey abundance. 


119

120 

EFFECT OF ROADS ON HABITAT USE BY COUGARS 

DOROTHY M. FECSKE, Department of Wildlife and Fisheries Sciences, South Dakota State 

University, Brookings, SD 57007, USA, email: gdf@rapidnet.com 

JONATHAN A. JENKS, Department of Wildlife and Fisheries Sciences, South Dakota State 

University, Brookings, SD 57007, USA, email: Jonathan_Jenks@sdstate.edu 

FREDERICK G. LINDZEY, USGS Biological Resources Division, Wyoming Cooperative Fish 

and Wildlife Research Unit, University Station, Laramie, WY 82071, USA, email: 

Flindzey@uwyo.edu 

STEVEN L. GRIFFIN, South Dakota Department of Game, Fish and Parks, 3305 W. South 

Street, Rapid City, SD 57702, USA, email: Steve.Griffin@state.sd.us 

Abstract: We examined effect of roads on habitat use by cougars, Puma concolor, in the Black 

Hills, South Dakota. A total of 768 daytime locations of 12 radio-collared cougars were 

obtained during weekly flights (1999 - 2001) using aerial telemetry techniques. Locations were 

incorporated into a geographic information system (GIS) of roads (Class 1, 2, 3, and 4). We 

tested the null hypotheses that cougars select habitat at random distances to roads and at random 

road densities and cougar use of habitat near roads did not differ with respect to road class, sex, 

age class, and habitat quality (based on a ranked cougar habitat-relation model). We examined 

use of habitat near roads for an adult female cougar fitted with a Global Position System (GPS) 

collar during crepuscular, diurnal, and nocturnal periods. Also, we identified road classes where 

cougar snow tracks were located and cougar/vehicle collisions occurred. During daylight hours, 

cougars avoided habitat near Class 3 roads (P < 0.001), the predominant road class in the Black 

Hills. However, on occasions where cougars were located near Class 3 roads, high quality 

habitat was selected. Cougars in the 5 to 6-year age class were located farther from Class 1 roads 

than younger animals (P < 0.0001). Females in the 1 to 2-year age class were located closer to 

Class 1 and Class 2 roads than older females (P < 0.0001). Females in 5 to 6 and 7 to 8-year age 

classes were located closer to Class 4 roads (P = 0.0047) than younger females. Road densities 

(km road/km 2 ) in annual home ranges of male cougars did not differ (P = 0.5000) from road 

densities throughout the Black Hills study area but densities in annual ranges of females were 

greater (P = 0.0078) than those of the study area. Cougars in the Black Hills have adapted to a 

heavily roaded landscape but presence of roads is impacting cougar use of habitat and survival. 

We suggest use of habitat near Class 3 roads by cougars would increase if roads were closed or 

had limited access, and if thinned ponderosa pine stands adjacent to Class 3 and 4 roads were 

managed for understory vegetation. 


ECOLOGY OF SYMPATRIC PUMAS AND JAGUARS IN NORTHWESTERN 

MEXICO 

CARLOS A. LOPEZ GONZALEZ, Universidad Autonoma de Queretaro, Cerro de las 

Campanas s/n, Mexico, email: Cats4mex@aol.com 

SAMIA E. CARRILLO PERCASTEGUI, Northern Jaguar Project, 2114 W. Grant Rd. #121, 

Tucson, AZ 85745, USA, email: Cats4mex@aol.com 

Abstract: Pumas (Puma concolor) are usually considered subordinate species where jaguars 

(Panthera onca) are present. Most of the current information on resource partitioning by these 

two species comes from tropical sites. Out study area is located in the limits of the tropical realm 

and consequently could be characterized as puma habitat. Our objectives were to describe the 

ecology of both large felids in an area located 135 s. of the United States border, in the state of 

Sonora, Mexico. From July 1999 to December 2002 using a suite of methodologies (cameratraps, 

radio-telemetry, scat, track and prey surveys), we surveyed an area ≈1000 km². The study 

area is a matrix of oak-woodland, tropical thornscrub, and upper sonoran desert; ranging in 

elevation from 200 to 1200 m. The main economic activity within the region is ranching. We 

determined through radio-telemetry a density of 3 pumas/100 km², and through camera-trap 

surveys a density of 1.4±0.4 jaguars/100 km². Camera-trap capture rates are three times higher 

for pumas than jaguars. Both species are feeding on white-tailed deer and to a lesser extent on 

livestock. Pumas are a cathemeral species whereas jaguars are nocturnal-crepuscular. Jaguars are 

using oak woodlands more than expected by chance, where pumas are using habitats according 

to availability. The number of pumas present may be an artifact of less prosecution by cowboys 

(only 1 puma killed since 1999), where jaguars are constantly prosecuted as they are perceived as 

liable of most livestock depredations (22 jaguars killed since 1999). During 2002 we began a 

program to help local ranchers on maintaining infrastructure, and apparent result has been less 

pressure on the jaguar population within the area. 


121

122 

COUGAR ECOLOGY AND COUGAR-WOLF INTERACTIONS IN YELLOWSTONE 

NATIONAL PARK: A GUILD APPROCH TO LARGE CARNIVORE CONSERVATION 

TONI K. RUTH, Associate Conservation Scientist, Wildlife Conservation Society, 2023 Stadium 

Dr. Suite 1A, Bozeman, MT 59030, USA, email: truth@montanadsl.net 

POLLY C. BUOTTE, Research Assistant, Wildlife Conservation Society, 2023 Stadium Dr. 

Suite 1A, Bozeman, MT 59030, USA, email: polly_thornton@hotmail.com 

HOWARD B. QUIGLEY, Beringia South, 3610 W. Broadwater, Suite 111, Bozeman, MT 

59715, USA 

MAURICE G. HORNOCKER, Senior Scientist, Wildlife Conservation Society, 2023 Stadium 

Dr. Suite 1A, Bozeman, MT 59030, USA 

Abstract: Successful restoration of large carnivores in the Northern Rockies and the concomitant 

increase in carnivore abundance and distribution will challenge humans as human development 

increases throughout the West. Presently, there is little understanding of how 

reintroduction/reestablishment of endangered large carnivores (wolves and grizzly bears) may 

affect the population characteristics, distribution, and behavior of other large carnivore 

populations, such as cougars. If restored wolves limit cougar populations in number or 

distribution, this limitation may have synergistic effects with current relaxation of cougar 

hunting regulations and rapid development. An added stress such as low prey availability (e.g. 

caused by hard winter or disease) could further impact populations. Understanding competitive 

relationships between large carnivores and the role that habitat and prey availability play is 

paramount to predicting and preparing for changes in the Greater Yellowstone region. In order 

to assess population-level effects of wolf (Canis lupus) reestablishment on cougars (Puma 

concolor) in and near Yellowstone National Park (YNP), we initiated a Phase II study of YNP 

cougars in 1998. The study is designed to examine the characteristics of the cougar population 

including: sex and age structure, density, reproductive and survival rates, dispersal and 

recruitment events, rate of predation on prey, and spatial and temporal movements. These 

parameters will be compared with analogous estimates made prior to the wolf restoration event 

in 1995 (Phase I data, Murphy 1998) and similar parameters documented for the wolf population 

to assess competition and resource partitioning between the two species. During 1998-2002, 56 

cougars were captured in and adjacent to areas used by 35-88 wolves within 3-5 wolf packs on 

the Northern Yellowstone Study Area, Montana and Wyoming. A sample of 3 to 10 radiocollared 

wolves was maintained within each wolf pack by the Yellowstone Wolf Restoration 

program. In this paper we summarize current research findings relative to cougar population 

changes pre- and post-wolf reintroduction, species interactions, and discuss future study 

direction. 


EVALUATION OF HABITAT FACTORS THAT AFFECT THE ABUNDANCE OF 

PUMAS IN THE CHIHUAHUAN DESERT 

JOEL LOREDO SALAZAR, Instituto de Ecología, A.C. Apdo. Postal 63. CP 91070 Jalapa, Ver. 

México, email: loredosj@ecologia.edu.mx 

LUCINA HERNÁNDEZ, Instituto de Ecología, A.C., Apartado Postal 632, 34100 Durango, 

Dgo., México, email: lucina@sequia.edu.mx 

JOHN W. LAUNDRÉ, Instituto de Ecología, A.C., Apartado Postal 632, 34100 Durango, Dgo., 

México, email: launjohn@prodigy.net.mx 

Abstract: Pumas originally occupied all of Mexico but their current status is not well known. 

This is especially true in the Chihuahuan desert of Northern Mexico. To manage this species in 

this area, it is important to have some estimation of their status. To evaluate the status of pumas 

in this area we need to first identify factors that may contribute to their rarity or abundance. 

Such factors can be placed into three separate but related groups: habitat quality, prey 

abundance, and human impacts. To evaluate these various factors, we selected two mountain 

ranges in the northern Chihuahuan desert where previous work indicated differences in the 

relative abundance of pumas. The area of low puma abundance was El Cuervo near Aldama, 

Chihuahua and the area of high abundance was Sierra Rica in the Canyon de Santa Elena 

protected area. In the field we estimated habitat quality by measuring shrub density, cover, and 

height. We also estimated prey (wild and domestic) abundance by counting fecal groups along 

random transects. With the use of GIS technology we assessed human impacts by determining 

the number of roads, number and size of towns, and overall density of humans in a 20 km radius 

around each range. Our results indicate that habitat quality was similar between the two areas. 

However, wild and domestic prey was higher in Santa Elena and all measurements of human 

impact were higher in El Cuervo. We concluded that habitat quality was not a factor 

contributing to relative puma abundance. However, the increased presence of and access by 

humans in El Cuervo is the main contributing factor via illegal hunting of pumas and their prey. 

Future work will test this hypothesis in other areas of the Chihuahuan desert. If this hypothesis 

is supported, it indicates that conservation efforts of pumas in Northern Mexico need center on 

environmental education rather than habitat protection/restoration. 


123

124 

ARE PUMAS OPPORTUNISTIC SCAVENGERS? 

JIM W. BAUER, University of California-Davis, Wildlife Health Center, Southern California 

Puma Project Field Station, P.O. Box 1203, Julian, CA 92036, USA, email: 

jwbauer@uia.net 

KENNETH A. LOGAN, University of California-Davis, Wildlife Health Center, Southern 

California Puma Project Field Station, P.O. Box 1114, Julian, CA 92036, USA, email: 

klogan2@mindspring.com 

LINDA L. SWEANOR, University of California-Davis, Wildlife Health Center, Southern 

California Puma Project Field Station, P.O. Box 1114, Julian, CA 92036, USA, email: 

lsweanor@mindspring.com 

WALTER M. BOYCE, University of California-Davis, Wildlife Health Center, One Shields 

Avenue, Davis, CA 95616, USA, email: wmboyce@ucdavis.edu 

Abstract: We examined scavenging on mule deer (Odocoileus hemionus) carcasses by pumas 

(Puma concolor) in the Peninsular Ranges of Southern California. Between 23 January 2001 

and 21 November 2002, a total of 42 deer carcasses from road kills, depredation permits, and 

euthanized deer were used to determine scavenging events. Seventeen of 42 deer carcasses 

(40.5%) were scavenged by 7 to 10 different pumas. Two of the scavenging pumas (males) were 

previously telemetered, while 4 pumas (3 male, 1 female) were captured and instrumented at the 

scavenging site. Telemetered pumas ranged in age from 11 months to 9 years. Deer carcasses 

were found and scavenged by pumas within 1 to 14 days, when carcass conditions ranged from 

fresh to rotting and maggot infested. Pumas treated scavenged carcasses as they would their own 

kills, dragging untethered carcasses to preferred sites and caching, as well as depositing scats and 

making scrapes in the area. However, pumas did not always attempt to cache tethered carcasses. 

During the course of our fieldwork we also discovered that one telemetered puma was repeatedly 

visiting a domestic livestock graveyard and scavenging on surface-discarded horse and cattle 

carcasses. While pumas are known to be opportunistic predators, our results would suggest that 

they are opportunistic scavengers as well. Due to pumas’ propensity to scavenge, it is likely that 

some perceived puma kills may in fact be scavenging events. Frequent monitoring and timely 

field investigation of mortality signals detected from telemetered prey species will help 

investigators identify those events. Scavenging behavior should be considered when evaluating 

or predicting the effects of puma predation on prey species. 


COUGAR-INDUCED INDIRECT EFFECTS: DOES THE RISK OF PREDATION 

INFLUENCE UNUGULATE FORAGING BEHAVIOR ON THE NATIONAL BISON 

RANGE? 

DAVID M. CHOATE, Ph.D. candidate, Department of Biological Sciences, University of Notre 

Dame, 107 Galvin Life Science Center, Notre Dame, IN 46556, USA; and, Department 

of Forestry, Range & Wildlife, Utah State University, Logan, UT 84322, USA, email: 

dchoate@nd.edu 

GARY E. BELOVSKY, Professor, Department of Biological Sciences, University of Notre 

Dame, 107 Galvin Life Science Center, Notre Dame, IN 46556, USA, email: 

Gary.E.Belovsky.1@nd.edu 

MICHAEL L. WOLFE, Professor, Department of Forestry, Range & Wildlife, Utah State 

University, Logan, UT 84322, USA, email: mlwolfe@cc.usu.edu 

Abstract: Ecologists have long debated whether predators (“top-down”) or nutrients/food 

(“bottom-up”) limit prey populations. Evidence supporting the importance of predation is 

frequently based on the number of prey killed by predators – a direct effect. By examining only 

this direct effect many predation studies fail to consider behavioral changes arising from the risk 

of predation - indirect effects. Furthermore, these behavioral indirect effects can be more 

important than the direct effect of predator-caused mortality, influencing both top-down and 

bottom-up processes. In this study we capitalize on a “natural experiment” on a suite of large 

mammalian herbivores, in a system (National Bison Range, MT) where the behavior and 

population dynamics of ungulate prey species (whitetail deer, Odocoileus virginianus; mule deer, 

O. hemionus; elk, Cervus elaphus) can be compared before and after an increase in risk of 

predation by cougar (Puma concolor). We present preliminary data demonstrating that cougars 

can influence several aspects of prey behavior. With an increase in predation risk, mule deer and 

elk total daily activity time has declined by 35.9% and 31.8% (P

126 

COUGAR PREDATION ON PREY IN YELLOWSTONE NATIONAL PARK: A 

PRELIMINARY COMPARISON PRE- AND POST-WOLF REESTABLISHMENT 

TONI K. RUTH, Wildlife Conservation Society, 2023 Stadium Dr. Suite 1A, Bozeman, MT 

59030, USA, email: truth@montanadsl.net 

POLLY C. BUOTTE, Wildlife Conservation Society, 2023 Stadium Dr. Suite 1A, Bozeman, MT 

59030, USA, email: polly_thornton@hotmail.com 

KERRY M. MURPHY, Yellowstone Center for Resources, P.O. Box 168, Yellowstone National 

Park, Mammoth, WY 89210, USA, email: kerry_murphy@nps.gov 

MAURICE G. HORNOCKER, Wildlife Conservation Society, 2023 Stadium Dr. Suite 1A, 

Bozeman, MT 59030, USA 

Abstract: On Yellowstone National Park’s Northern Range cougars and wolves rely on 

economically important prey species, particularly elk. Understanding how these large carnivores 

partition prey resources and their combined affect on prey is important for management and 

conservation of cougars, wolves, and ungulate species. As part of a cougar-wolf interactions 

study, we quantified predation rates and prey selection by cougars on Yellowstone’s northern 

range prior to (Phase I) and post wolf (Phase II) reestablishment. During Phase II, cougars spent 

an average of 3.7 days at kills and 4.4 days between each kill. The mean annual rate of cougar 

predation in Phase I was 9.4 (SD = 4.0; 95% CI = 7.8 to 11.0) days per ungulate kill, and 

10.9(SD = 8.5; 95% CI = 6.7 to 15.1) days per ungulate kill in Phase II. Rate of predation varied 

by cougar social class. When converted to biomass killed per day, cougars averaged 12.2 kg per 

day during Phase I and 12.9 kg per day during Phase II. We documented a total of 306 and 256 

positive and probable cougar kills during Phase I and Phase II, respectively. During Phase II, 

70% (n = 179) of cougar kills were elk, 17% (n = 43) were mule deer and 13% (n = 34) were 

other prey. During both Phase I and II more than 50% of cougar kills were elk calves, with cow 

elk making up the next largest category. During Phase I, cougar predation was neither a major 

source of mortality nor a significant factor limiting the numbers or growth rates of elk and mule 

deer populations in northern Yellowstone. Cougars present on the study area killed 2-3% of the 

elk and 3-5% of the mule deer estimated to be available during 5 years spanning the Phase I 

study. Simultaneous to our Phase II study, the Yellowstone Wolf Project quantifies wolf 

predation rates and prey selection. Cougars killed proportionally more elk calves and fewer bull 

elk than wolves between 1998 and 2002. We are continuing our data collection and analyses and 

plan to: 1) compare cougar and wolf per capita rate of predation, 2) contrast femur marrow fat 

content of cougar and wolf kills, by season killed and prey age, and 3) compare yearly off-take 

of elk and mule deer by cougars and wolves. Cougar per capita predation rate averaged across 

social classes was 0.06 kills/cougar/day. When kittens were included with maternal females, that 

group had the lowest predation rate of 0.01. Without including kittens, maternal females 

averaged 0.15. Subadult males had an equally high rate of 0.15. Wolf predation ranged from 

0.03 to 0.078 kills per wolf per day (Smith et al., In Press). 


FOUR DECADES OF COUGAR-UNGULATE RELATIONSHIPS IN THE CENTRAL 

IDAHO WILDERNESS 

HOLLY A. AKENSON, University of Idaho, Taylor Ranch Field Station, HC 83 Box 8070, 

Cascade, ID 83611, USA, e-mail: tayranch@direcpc.com 

JAMES J. AKENSON, University of Idaho, Taylor Ranch Field Station, HC 83 Box 8070, 

Cascade, ID 83611, USA; e-mail: tayranch@direcpc.com 

HOWARD B. QUIGLEY, Beringia South, 2023 Stadium Drive, Suite 1A, Bozeman, MT 

59715, USA 

MAURICE G. HORNOCKER, Wildlife Conservation Society, 2023 Stadium Drive, Suite 1A, 

Bozeman, MT 59715, USA 

Abstract: Research conducted on cougars (Puma concolor) in the Big Creek drainage in each of 

the last four decades has enhanced the understanding of the dynamic nature of cougar – ungulate 

relationships. In 1964, Maurice Hornocker initiated his benchmark research on this cougar 

population and assessed the role of cougar predation in regulating ungulate populations. Each 

study that followed has had different objectives, yet, combined these projects provide a rare 

continuum of ecological information on the dynamics of cougar – prey relationships. This 

cougar population has been influenced by significant environmental changes over the last 40 

years. The ungulate prey base has fluctuated, but generally elk numbers have increased and deer 

have decreased. Total ungulate biomass was similar in the 1960’s and 1980’s, but was 12% 

lower in the study just completed. The dynamics of carnivore competition, both inter-specific 

and intra-specific, has changed since introduced wolves recolonized the drainage in the 1990s. A 

large-scale forest fire 2 years ago drastically altered winter and summer ranges and affected 

predator – prey relationships. We compared cougar population size, structure, reproduction, and 

mortality factors; prey selection during 3 time periods; and evaluated pre and post-fire data in the 

recent study. The estimated resident cougar population was 9 adults during the first 2 studies in 

the 1960’s and early 1970’s. The resident population grew to an estimated 13 adults in the mid- 

1980’s, but dropped to 10 individuals by 2000, and down to 6 resident cougars by 2002. The 

population increase during the 1980’s was in the adult female segment and it corresponded with 

an increasing elk population. The current low population is a result of a decreasing elk 

population, ungulate displacement from fire, increased hunter harvest of cougars, increased 

intraspecific strife, and competition with wolves for the same prey base. Cougars selected for 

elk rather than mule deer during the first study, but killed elk in proportion to their relative 

abundance during the study in the 1980’s and recent study (2000). Historical perspectives from 

pioneer diaries indicate similar cougar population numbers. In 1888 a bounty hunter removed 12 

cougars from the drainage, then ten years later a different cougar hunter noted trapping and 

poisoning 12 individuals on Big Creek. Archeological evidence, old newspaper articles and 

diaries, and early agency field notes are all integrated into this discussion of long-term predator - 

prey relationships. The lengthy record of information on predator and prey populations in the 

Big Creek drainage arguably makes this cougar population the best understood in North 

America. 


127

128 

COUGAR TOTAL PREDATION RESPONSE TO DIFFERING PREY DENSITIES: A 

PROPOSED EXPERIMENT TO TEST THE APPARENT COMPETITION 

HYPOTHESIS 

HUGH ROBINSON, Large Carnivore Conservation Lab, Department of Natural Resource 

Sciences, Washington State University, PO Box 646410, Pullman, WA 99614-6410, 

USA, email: hsrobins@wsunix.wsu.edu 

ROBERT WIELGUS, Large Carnivore Conservation Lab, Department of Natural Resource 


USA, email: wielgus@wsu.edu 

HILARY CRUICKSHANK, Large Carnivore Conservation Lab, Department of Natural 

Resource Sciences, Washington State University, PO Box 646410, Pullman, WA 99614- 

6410, USA, email: hcruicks@mail.wsu.edu 

CATHERINE LAMBERT, Large Carnivore Conservation Lab, Department of Natural Resource 


USA, email: lambertcath@wsu.edu 

Abstract: Mule deer populations throughout the west are declining whereas white-tailed deer 

populations are increasing. We compared abundance, fetal rate, recruitment rate, and causespecific 

adult (≥1 yr. old) mortality rates of sympatric mule and white-tailed deer in south-central 

British Columbia to assess the population growth of each species. White-tailed deer were three 

times as abundant (908±152) as mule deer (336±122) (± 1SE). Fetal rates of white-tailed deer 

(1.83) were similar to mule deer (1.78) (t = 0.15, df = 13, P = 0.44) as was recruitment of whitetailed 

deer (56 fawns:100 does) and mule deer (38 fawns:100 does) (χ 2 = 0.91, df = 1, P=0.34). 

Annual adult white-tailed deer survival (SWT = 0.81) was significantly higher (z = 1.32, df = 1, P 

= 0.09) than mule deer survival (SMD = 0.72). The main source of mortality in both populations 

was cougar predation. The lower survival rate of mule deer could be directly linked to a higher 

predation rate (0.17) compared to white-tailed deer (0.09) (z = 1.57, df = 1, P = 0.06). The finite 

growth rate (λ) of mule deer was 0.88 and 1.02 for white-tailed deer. We suggest that the 

disparate survival and predation rates are caused by apparent competition between the two deer 

species, facilitated through a shared predator; cougar. The apparent competition hypothesis 

predicts that as alternate prey (white-tailed deer) densities increase, so do densities of predators, 

resulting in increased incidental predation on sympatric native prey (mule deer). Apparent 

competition can result in population declines and even extirpation of native prey in some cases. 

Such a phenomenon may account for declines of mule deer throughout the arid and semi-arid 

West where irrigation agriculture is practiced. We are in year two of a proposed five-year study. 

We will test the apparent competition hypothesis by conducting a controlled, replicated “press” 

experiment in 2 treatment and 2 control areas in North-eastern Washington by reducing densities 

of white-tailed deer and observing any changes in cougar predation on mule deer. Washington 

Fish and Wildlife personnel using annual aerial surveys and/or other trend indices will monitor 

deer densities. Predation rates and population growth rates of deer will be determined using radio 

telemetry. Changes in cougar functional (kills/unit time), aggregative (cougars/unit area), 

numerical (offspring/cougar), and total (predation rate) responses on deer will also be monitored 

using radio telemetry. Results will be used to determine the effect of increased white-tailed 

densities on cougar predation of mule deer. 


CHARACTERISTICS OF COUGAR HARVEST WITH AND WITHOUT THE USE OF 

DOGS 

DONALD A. MARTORELLO, Washington Department of Fish and Wildlife, 600 Capitol Way 

North, Olympia, WA 98501-1091, USA, email: martodam@dfw.wa.gov 

RICHARD A. BEAUSOLEIL, Washington Department of Fish and Wildlife, 3515 State 

Highway 97A, Wenatchee, WA 98801, USA, email: beausrab@dfw.wa.gov 

Abstract: Prior to 1996, dogs were used to harvest the majority (99%) of cougar during recreational hunting seasons 

in Washington State. However, in 1996 Voter Initiative 655 banned the use of dogs to aid in the harvest of cougar. 

As a result, harvest methods shifted to spot and stalk, predator calling, and incidental encounters between deer and 

elk hunters and cougar. We examined the sex and age structure of harvested cougar and compared these data 

between seasons with (selective harvest) and without the use of dogs (non-selective harvest). We detected a 

significant increase in percent female cougars in the total harvest, from 42% to 59% during selective versus nonselective 

seasons (T = -7.85, P < 0.0001). We also found that non-selective harvest seasons had significantly more 

2 

2 

juvenile male ( χ = 98.1790, d.f. = 10, P < 0.0001) and female ( χ = 66.5116, d.f. = 10, P < 0.0001) cougars 

compared to selective seasons. We then used program RISKMAN to evaluate the potential impacts to population 

growth (finite rate of increase) from changes we observed in harvest vulnerability of specific sex and age classes. 

Our sensitivity analysis suggests that changes in female adult and cub survival are the most influential parameters to 

population growth and the increased harvest of female cougars in non-selective harvest methods decreased the finite 

rate of increase by about 0.01–0.02. Harvest methods that increase the relative harvest vulnerability of these cohorts 

have a greater potential for impacting population growth. In Washington State, the current level of cougar harvest 

and increased vulnerability of females and juvenile cougar have likely increased the risk of impacting population 

growth. 

129 


Key Words: Voter Initiative 655, selective harvest, non-selective harvest, Washington, cougar, Puma concolor, 

RISKMAN 

INTRODUCTION 

In Washington State, cougar (Puma 

concolor) hunting regulations and harvest 

levels have changed dramatically during the 

last decade (Table 1). Pivotal to these 

changes was Voter Initiative 655 (I-655), 

which banned the use of dogs for hunting 

cougar in Washington in 1996. Prior to I- 

655, hunters used dogs to take about 99% of 

the harvested cougar. Recognizing that 

using dogs was more effective than spotand-stalk 

or predator calling methods to 

harvest cougar, Washington Department of 

Fish and Wildlife (WDFW) instituted 

several changes in an effort to maintain 

harvest levels similar to pre-initiative 

seasons; including lengthening the season 

from about 86 to 227 days, increasing the 

annual bag limit from 1 to 2 cougars/hunter, 

and decreasing the cost of a cougar transport 

tag from $24 to $10. 

Both prior to and since I-655, wildlife 

managers in Washington used harvest 

characteristics to assess the status of the 

living cougar population. Managers 

monitored trends in total harvest, harvest 

success, percent females in the harvest, and 

median ages of harvested males and 

females. Using harvest information alone to 

assess a living population is fraught with 

problems. The value of harvest information 

becomes even more suspect when the 

harvest methods change, because the 

relationship between trend data and the

130 CHARACTERISTICS OF COUGAR HARVEST · Martorello and Beausoleil 

Table 1. Recreational cougar hunting seasons in Washington, 1990-2001. 

Year Season Dates Days Harvest Restrictions Selectivity 

1990-1991 Nov. 21 – Jan. 15 57 102 Permit only season Dogs allowed 

1991-1992 Nov. 27 – Jan. 15 50 120 Permit only season Dogs allowed 

1992-1993 Oct. 17 – Jan. 31 107 140 Permit only season Dogs allowed 




1996-1997 Oct. 12 – Mar. 15 155 178 General season No dogs allowed 

1997-1998 Aug. 1 – Mar. 15 227 132 General season No dogs allowed 




living population also may have changed. 

As a result, trend information based on 

harvest data may be of limited value in 

Washington because of the restriction on 

using dogs to hunt cougar. 

Little information is known about how 

restrictions on using dogs to hunt cougar 

impacts harvest, trends based on harvest 

data, or cougar populations. To that end, 

our objectives are to: 1) determine if there 

are differences in cougar harvest 

characteristics (i.e., total harvest, percent 

female cougar in the harvest, and age 

structure of harvest cougar) during seasons 

with and without the use of dogs, and 2) 

identify potential impacts to the cougar 

population given changes in season structure 

and harvest levels in Washington between 

1990-2001. 

METHODS 

We obtained cougar harvest data from 

1990 to 2001 through a mandatory harvest 

reporting system implemented by WDFW. 

Successful hunters were required to report 

their harvested cougar and present the hide 

and skull to WDFW, where Agency staff 

collected a tooth sample, and documented 

sex, kill location, kill date, and kill type 

(depredation, recreational, public safety 

cougar removal, or other). We determined 


ages for 79% of the harvested cougar using 

cementum annuli analysis. 

We classified harvest data from 1990- 

1995 as “selective” and from 1996-2001 as 

“non-selective” to compare harvest 

characteristics between periods when dogs 

were legal to years when they were not, 

respectively. We used a T-test to determine 

if cougar harvest (recreational only) and 

percent females in the harvest (recreational 

only) differed between selective versus nonselective 

periods. To compare harvest age 

structures between selective and nonselective 

periods, we generated a mean age 

structure for each sex and each period by 

pooling cougar management units (CMUs) 

(Washington is divided into 9 administrative 

CMUs) and years. We then used a chisquare 

test to determine if mean age 

structures for each sex differed between 

selective versus non-selective periods. We 

considered all tests significant at α ≤ 0. 

05 . 

We used program RISKMAN to assess 

potential impacts to the cougar population 

from changes in harvest methods and rates 

(Taylor et al. 2002). We incorporated 

parameter estimates from Logan and 

Sweanor (2001) and Spencer et al. (2001) 

(Table 2). For initial population size we 

multiplied the amount of suitable cougar 

habitat in Washington (88,497 km 2 ) by the

CHARACTERISTICS OF COUGAR HARVEST · Martorello and Beausoleil 131 

Table 2. Parameter inputs for program RISKMAN simulations in Washington. 

Parameter Value Source 

Recruitment Probability of 1 cub 0.00 Logan and Sweanor 2001 

Probability of 2 cubs 0.25 Logan and Sweanor 2001 



Mean litter size 3.01 Logan and Sweanor 2001 

Proportion of females with litters 0.80 Logan and Sweanor 2001 

Proportion of males at birth 0.50 Logan and Sweanor 2001 

Survival Male cubs (age 0) 0.67 Logan and Sweanor 2001 

Female cubs (age 0) 0.67 Logan and Sweanor 2001 

Male yearling (age 1-2) 0.64 Logan and Sweanor 2001 

Female yearlings (age 1-2) 0.88 Spencer et al. 2001 

Male adults (age 3-12) 0.91 Logan and Sweanor 2001 

Female adults (age 3-12) 0.82 Logan and Sweanor 2001 

Litter survival rate 0.93 Logan and Sweanor 2001 

Population size 4159 WDFW a , unpublished data 

a 

Washington Department of Fish and Wildlife 

highest cougar density reported in the 

literature (4.7 cougars/100 km 2 ) (Ross and 

Jalkotzy 1992). Each of the parameters 

values, including density, are at the high end 

in terms of the range reported in the 

literature. We chose to model a population 

with high productivity, survival, and density 

so our analysis would reflect how harvest 

strategies and rates might impact even the 

most robust cougar population. Therefore, 

potential impacts to a more realistic 

population would be at least as great, if not 

more, than those to our modeled population. 

We used the age structure from cougars 


harvested between 1997-1999 to develop the 

input age distribution for the model. We 

selected these years because they probably 

better reflect the standing population than 

years when dogs were legal or years when 

potential impacts may begin to be apparent. 

From the average age structure we generated 

a stable age distribution and used this as the 

age distribution in the model. 

To assess the impacts of selective versus 

non-selective harvest methods, we 

conducted two simulations; one where the 

strata specific harvest vulnerabilities were 

set to levels observed during the selective 

Table 3. Vulnerability inputs for selective and non-selective model simulations in Washington. 

Relative vulnerability 

Parameter Age class Selective Non-selective 

Cubs (age 0) 0.000 1.0 

Males 

Females without offspring 

Juveniles (age 1-2) 0.118 1.0 

Adults (age 3-12) 0.464 1.0 

Cubs (age 0) 0.001 1.0 

Juveniles (age 1-2) 0.109 1.0 

Adults (age 3-12) 0.277 1.0 

Females with offspring Adults (age 3-12) 0.031 1.0


period (1990-1995) and one where the 

vulnerabilities were constant across strata, 

thereby mimicking a non-selective method 

(1996-2001) (Table 3). We repeated each 

simulation for harvest rates ranging from 0% 

(no harvest) to 14% of the censused 

population. All simulations were 

deterministic and the realized finite rate of 

increase with no harvest was 1.062. 

RESULTS 

Mean harvest increased from 157 during 

the selective period to 199 in the nonselective 

period; however the increase was 

not statistically significant (T = -1.26, P = 

0.2368). We detected a significant increase 

in percent females in the harvest (T = -7.85, 

P < 0.0001). Percent females in the harvest 

increased from 42 to 59% during selective 

versus non-selective periods, respectively 

(Figure 1). We detected a significant 

difference in male age distributions for 

2 

selective versus non-selective periods ( χ = 

98.1790, d.f. = 10, P < 0.0001). On a 

statewide level, there were a greater 

proportion of younger males in the harvest 

during the non-selective period. Although 

less pronounced, there also were a lower 

proportion of adult males in the non- 

Recreational harvest 

300 

250 

200 

150 

100 

50 

0 

1990 

1991 

1992 

1993 

1994 

1995 

1996 

1997 

1998 

1999 

2000 

2001 

Year 

Recreational harvest % Female in harvest 

0.7 

0.6 

0.5 

0.4 

0.3 

0.2 

0.1 

0 

Figure 1. Recreational cougar harvest and 

percent female in the harvest during selective 

(1990-1995) and non-selective (1996-2001) 

years, Washington, 1990-2001. 

% Female in harvest 


selective period (Figure 2). We detected a 

significant difference in female age 

distributions for selective versus non- 

2 

selective periods ( χ = 66.5116, d.f. = 10, P 

< 0.0001). On a statewide level, there also 

were a greater proportion of younger 

females in the harvest during the nonselective 

period (Figure 2). 

Using model simulations, we found that 

changing from a selective harvest to a nonselective 

harvest (with all other parameters 

held constant) decreased the population’s 

finite rate of increase by about 0.01–0.02 

annually (Figure 3). This is roughly 

equivalent to about a 1.5% increase in 

harvest rate (proportion of population 

harvested). 

Frequency 

Frequency 

25 

20 

15 

10 

5 

0 

30 

25 

20 

15 

10 

5 

0 

A. 

0 1 2 3 4 5 6 7 8 9 10 11 12 13 16 17 

Age 

B. 

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 

Age 

1990-1995 1996-2001 

Figure 2. Age structures of harvested male 

(A) and female (B) cougar during selective 

(1990-1995) and non-selective (1996-2001) 

periods, Washington, 1990-2001.

Finite rate of increase 

1.08 

1.06 

1.04 

1.02 

1.00 

0.98 

0.96 

0.94 

0.92 

0.90 

Non-selective 

0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 

Harvest rate 

Figure 3. Relationship between harvest rate 

and finite rate of increase for selective and 

non-selective harvest strategies. 

DISCUSSION 

In Washington, when dogs were legal 

for hunting cougar, seasons primarily 

occurred from October to January. In 

contrast, when dogs were banned, season 

length increased and, more importantly, 

overlapped with deer and elk seasons. The 

reduced cougar tag and overlapping seasons 

made purchasing a cougar tag more 

attractive for deer or elk hunters, and 

licensed cougar hunters increased from less 

than 1,000 annually prior to I-655 to about 

58,000 post I-655. This in turn created a 

situation where the majority of the harvest 

was by deer and elk hunters that took a 

cougar incidentally during their deer or elk 


Selective 


hunt. We believe season timing, tag cost, 

and the large number of deer and elk hunters 

resulted in post I-655 harvest levels that 

were similar to pre I-655 levels. 

The proportional increase of females and 

juveniles in the harvest during years with 

and without dogs probably was correlated 

with the proportions of these cohorts in the 

standing population. That is, individual sex 

and age classes of cougar were probably 

taken relative to their availability. 

Deviations between sex and age specific 

proportions in the population versus harvest 

probably were influenced by cougar and 

hunter distributions, cougar behavioral 

patterns and home range sizes, and prey 

distribution. 

Our model simulations suggest that 

population growth is sensitive to female 

survival. The importance of female survival 

for population growth has been well 

documented (Clark 1999). A simple 

sensitivity analysis done by halving each 

parameter one-by-one and documented the 

percent decline in lambda further illustrates 

that female survival is the most influential 

parameter on lambda (Table 4). It therefore 

seems reasonable that changing from 

selective to non-selective harvest methods 

can impact a population’s growth rate. Our 

model, although crude, suggests that the 

Table 4. Percent decrease in λ when cougar parameter values are divided by two. Realized 

λ =1.062 with original parameter values (see Table 2 for parameter estimate sources) (adapted 

from Clark 1999). 

Actual Reduced 

Percent decrease 

Parameter 

estimate estimate λ 

in λ 

Female survival 0.82 0.41 0.73 31.3 

Female juvenile survival 0.88 0.44 0.84 20.9 

Cub survival 0.67 0.34 0.94 11.5 

Litter size 3.01 1.50 0.94 11.5 

Litter survival 0.93 0.47 0.96 9.6 

Prop. of females with litters 0.80 0.40 0.97 8.7 

Male adult survival 0.91 0.46 1.06 0.0 

Male juvenile survival 0.64 0.32 1.06 0.0


impact may be in the range of increasing 

lambda by 0.01–0.02. In more familiar 

terms, this is analogous to a 1.5% increase in 

harvest rate. At first glance this appears 

small, but in our example population a 1.5% 

increase in harvest rate equals a 38% 

increase in observed harvest level. So 

changing to a non-selective harvest method 

was biologically equivalent to increasing the 

harvest by about 38%. 

Of course, our model and the 

corresponding impacts to population growth 

assume that harvest is constant between 

selective and non-selective harvest methods. 

This is probably not a reasonable 

assumption unless season adjustments are 

made to mitigate the inefficiencies of boot 

hunting (e.g., spot and stalk, predator 

calling, and incidental take) for killing 

cougar. Without season changes, similar to 

those we discussed earlier, harvest would 

probably decline significantly without the 

use of dogs. The population growth rate 

would likely increase accordingly until 

density dependence occurred. 

We also assumed there was no 

immigration or emigration occurring. Logan 

and Sweanor (2001) found that immigration 

could have as large of an impact to 

population growth as reproduction. This 

suggests that immigration could potentially 

counter decreases in lambda that resulted 

from increased female harvest. The effect of 

immigration acting in this manor decreases 

as the perimeter-to-area ratio of the 

population boundary decreases. 

MANAGEMENT IMPLICATIONS 

Several factors probably influence 

whether changing from a selective to nonselective 

harvest method will cause a cougar 

population to decline. In addition to 

parameter estimates and standard errors, 

knowing the age structure and growth rate 

are essential for predicting how an actual 

population might respond to changes in 

harvest vulnerability (Caughley 1977). 


Unfortunately, collecting biological data on 

cougar population dynamics is difficult and 

costly because cougars are extremely 

secretive, difficult to count, have large home 

ranges, and occur at relatively low densities. 

As such, wildlife managers often use harvest 

information as a surrogate to data on the 

living population. The risk of using harvest 

information is more acceptable when the 

majority of the harvest is males, because 

male survival has a relatively small impact 

on population growth. In contrast, the risk 

of using harvest data to guide management 

decisions is higher when the majority of the 

harvest is females, because female survival 

has a greater impact on population growth. 

ACKNOWLEDGMENTS 

We thank J. Beecham and K. Logan for 

reviewing earlier versions of our model 

simulations. This study received funding 

from Federal Aid in Wildlife Restoration, 

Project W-97-R. 


CAUGHLEY, G. 1977. Analysis of vertebrate 

populations. John Wiley and Sons, New 

York, New York, USA. 

CLARK, J.D. 1999. Black bear population 

dynamics in the Southeast: some new 

perspectives on some old problems. 

Eastern Black Bear Workshop 

Proceedings 15:97-115. 


Desert puma: evolutionary ecology and 

conservation of an enduring carnivore. 

Island Press. Washington D. C., USA. 

ROSS, P.I., AND M.G. JALKOTZY. 1992. 

Characteristics of a hunted population of 

cougars in southwestern Alberta. 


426. 

SPENCER, R.D., D.J. PIERCE, G.A. 

SCHIRATO, K.R. DIXON, AND C.B. 

RICHARDS. 2001. Mountain lion home 

range, dispersal, mortality and survival 

in the western Cascades Mountains of

Washington. Final Report. Washington 

Department of Fish and Wildlife, 

Olympia, Washington, USA. 



TAYLOR, M., M. OBBARD, B. POND, M. 

KUC, D. ABRAHAM. 2002. Riskman: 

user manual. The Queens Printer for 

Ontario, Ontario, Canada.

136 

DEFINING AND DELINEATING DE FACTO REFUGIA: A PRELIMIARY ANALYSIS 

OF THE SPATIAL DISTRIBUTION OF COUGAR HARVEST IN UTAH AND 

IMPLICATIONS FOR CONSERVATION 

DAVID C. STONER, Utah State University, Dept. of Forest, Range, and Wildlife Sciences, 

5230 Old Main Hill, Logan, UT 84322-5230, USA, email: dstoner@cc.usu.edu 

MICHAEL L. WOLFE, Utah State University, Dept. of Forest, Range, and Wildlife Sciences, 

5230 Old Main Hill, Logan, UT 84322-5230, USA, email: mlwolfe@cnr.usu.edu 

Abstract: Cougars (Puma concolor) in Utah are managed at two scales, often with differing 

objectives. The statewide population is managed for persistence and sustainable hunting 

opportunities, while at the finer scale of an individual management unit, a sub-population may be 

managed to accomplish density reductions, depending on local priorities. However, cougars 

have large and variable spatial requirements, and management unit boundaries may not coincide 

with actual demes. Compounding these constraints, the lack of cost-effective and reliable 

enumeration techniques increases the risk of inadvertent over-harvest. Current research suggests 

that for species difficult to enumerate, greater emphasis be placed on metapopulation-scale 

management in order to minimize the effects of uncertainties with respect to demography and 

dispersal behavior. Because harvest is the primary variable that managers can manipulate and 

measure, it is important to understand how recruitment patterns in minimally exploited 

populations may influence the persistence and recovery of heavily exploited populations. In this 

paper we discuss some of the factors that account for the spatial distribution of harvest and how 

this information can be used to develop management strategies in the absence of census data. 

We used 6 years of radio-telemetry data from a lightly exploited population in the Oquirrh 

Mountains of north-central Utah to quantify the effect of a small sanctuary (480 km²) on cougar 

survivorship, fecundity, and dispersal. We then mapped the locations of cougars harvested 

across the state from 1996-2001, and attempted to: (1) identify the factors that influence these 

patterns, and (2) determine the size and distribution of potential harvest sinks and de facto 

refugia in the state. Finally, we identified habitat patches on the periphery of the state that 

straddle management jurisdictions, representing areas of possible inter-state cooperation. We 

recommend that managers consider a metapopulation perspective and attempt to distribute 

harvest pressure in a spatially and ecologically relevant manner. In the absence of large (2400 

km²), contiguous refugia, small sanctuaries adjacent to areas of high exploitation may be mapped 

and utilized as a deterrent against potential over-exploitation. 


MONITORING CHANGES IN COUGAR SEX/AGE STRUCTURE WITH CHANGES IN 

ABUNDANCE AS AN INDEX TO POPULATION TREND 

CHUCK R. ANDERSON, JR., Wyoming Cooperative Fish and Wildlife Research Unit, Box 

3166, University Station, Laramie, WY 82071, USA, email: cander@uwyo.edu 


University Station, Laramie, WY 82071 USA, email: flindzey@uwyo.edu 

Abstract: Cougar (Puma concolor) management has traditionally been plagued by the inability to 

identify population trends for adequate assessment of management strategies. Monitoring 

changes in harvest sex-age structure as an index to population trends appears useful in tracking 

black bear populations and may be applicable to cougar management, especially given their strict 

social structure and territorial behavior. We documented changes in cougar harvest structure 

(sex-age) through experimental population reduction and recovery to better understand the 

relationship between sex-age composition and population trend in exploited populations. The 

cougar population in the Snowy Range, southeast Wyoming, declined from 58 (90% CI = 36 to 

81) in the fall of 1998 to 20 (90% CI = 14 to 26) independent cougars (>1 year old) by the spring 

of 2000 following 2 years of increased exploitation (mean exploitation rate = 43%) and increased 

to 46 (90% CI = 33 to 60) by the spring of 2003 following 3 years of reduced harvest levels 

(mean exploitation rate = 18%). Pre-treatment harvest composition was 63% subadults (1.0-2.5 

years old), 24% adult males, and 14% adult females (2 seasons; n = 22). A reduction in subadult 

harvest, an initial increase followed by a reduction in adult male harvest, and a steady increase in 

adult female harvest was consistent with hypothesized harvest vulnerability for a declining 

population. Harvest composition was similar at high and low densities with light harvest, but the 

proportion of male subadults increased at low density as adult males removed during the 

treatment period (high harvest) were replaced. Examining cougar sex ratios (m:f) alone appears 

to be of limited utility for identifying population change. Including age class, however, provides 

a useful metric in monitoring cougar population trend. We feel this approach could be applied to 

adaptively manage cougar populations where adequate sex and age data are collected from 

harvested animals. 


137

138 

MANAGEMENT OF COUGARS (PUMA CONCOLOR) IN THE WESTERN UNITED 

STATES 

DEANNA DAWN, San Jose State University, Biology Department 

MICHEAL KUTILEK, San Jose State University, Biology Department 

RICK HOPKINS, Live Oak Associates, Inc. 

SELEHKA ANAND, San Jose State University, Biology Department 

STEVE TORRES, California Department of Fish and Game 

Abstract: In the U.S., cougar (Puma concolor) populations still exist in 13 western states. While 

sport hunting of cougars remains a management goal for 10 of these states, there is little 

information on how different hunting harvest strategies affect their biology. Both the rate of 

harvest and the percentage of females in the harvest affect population stability. Therefore, the 

purpose of this study was to examine the effect of different harvest strategies on the harvest rate 

and the percentage of females in the harvest. Annual hunting harvest records were requested 

from all 10 states and were summarized into a database for analysis. Harvest strategies that 

included female sub-quotas were associated with the lowest percentage of females removed, 

however they also had some of the highest annual rates of harvest. These results suggest that, for 

some states, management strategies used in regulating sport hunting may offer little protection 

against over-harvesting the population. 


DYNAMICS AND VIABILITY OF A COUGAR POPULATION IN THE PACIFIC 

NORTHWEST 

CATHERINE LAMBERT, Large Carnivore Conservation Laboratory, Department of Natural 

Resource Sciences, Washington State University, Pullman, WA 99164-6410, USA, 

email: cathlambert@mac.com 

ROBERT B. WIELGUS, Large Carnivore Conservation Laboratory, Department of Natural 


email: wielgus@wsu.edu 

HUGH S. ROBINSON, Large Carnivore Conservation Laboratory, Department of Natural 


email: hsrobins@wsunix.wsu.edu 

DONALD D. KATNIK, Large Carnivore Conservation Laboratory, Department of Natural 

Resource Sciences, Washington State University, Pullman, WA 99164-6410, USA 

HILARY CRUICKSHANK, Large Carnivore Conservation Laboratory, Department of Natural 


email: hcruicks@mail.wsu.edu 

ROSS CLARKE, Columbia Basin Fish & Wildlife compensation program, 103-133 Victoria St., 

Nelson, BC V1L 4K3, Canada 

Abstract: Cougar (Puma concolor) populations are believed to be at high density and increasing 

throughout western North America, especially in the Pacific Northwest, as evidenced by 

increasing cougars/humans encounters. Harvest rates have increased as a result. To test this 

hypothesis, we determined the density, fecundity, survival, and growth rate of a cougar 

population in northeastern Washington, northwestern Idaho, and southern British Columbia. 

From 1998 to 2003, 52 cougars were captured, radio-collared, and monitored. We recorded 

fecundity through den site investigation and snow tracking, and mortality by weekly telemetry. 

Survival rates were estimated for kittens (0-1 yr), yearlings (1-2 yr), and adult (2-12 yr) males 

and females. Average overall density was 1.09 cougars/100km 2 or 0.46 adults/100km 2 . We 

estimated litter size at 2.53 kittens, birth interval at 18 months, proportion of reproductively 

successful females at 0.75, and age of first reproduction at 30 months, for a maternity rate of 

0.63 male or female kitten/year/adult female. Average survival rate for all radio-collared 

cougars was 0.59, 0.77 for adult females, 0.44 for adult males, 0.37 for yearlings, and 0.57 for 

kittens. Hunting accounted for 92% of the mortalities of radio-collared cougars. Age- and sexspecific 

survival and fecundity were entered into a stochastic two-sex matrix model. We used 

computer simulations to determine the population stochastic growth rate and to assess its 

viability over 25 years. The annual stochastic growth rate of this population was λ = 0.80 

(95%CI = 0.11). Starting with a total initial abundance of 357, the median times to fall below a 

demographic collapse (N = 30 adults) and extirpation (N = 0) were 8.5 and 25.9 years. Our 

findings suggest that, contrary to popular belief, cougars in the Pacific Northwest are currently at 

low to moderate densities and are declining. Alternative hypotheses may account for the 

increased conflicts between cougars and humans in this area. 


139

140 

PROJECT CAT (COUGARS AND TEACHING): INTEGRATING SCIENCE, SCHOOLS 

AND COMMUNITY IN DEVELOPMENT PLANNING 

GARY M. KOEHLER, Wildlife Research Scientist, Washington Department of Fish and 

Wildlife, 600 Capitol Way N., Olympia, WA 98501 USA, email: koehlgmk@dfw.wa.gov 

EVELYN NELSON, Superintendent, Cle Elum-Roslyn School District, 2690 SR 903, Cle Elum, 

WA 98922, USA, email: Nelsone@cleelum.wednet.edu 

Abstract: Complaint reports of cougars (Puma concolor) venturing into urban areas, killing 

livestock and pets, and threatening humans have increased to more than 1,000 reports filed 

annually in Washington; where in the past 5 years cougars have mauled 2 children. Increased 

reports are coupled with human population increases of over 1 million in the past decade and an 

annual loss to development of over 28,000 ha of land. The rural Cle Elum-Roslyn community is 

experiencing similar growth and development with >5,000 ha of development and over 1,400 

new homes planned, but presently with few complaints of cougars. The winter of 2002-2003 

marked the beginning of the 2 nd year of an 8-year scientific investigation on cougar and ecology 

by the Washington Department of Fish and Wildlife and data collection and analysis by teachers 

and students at Cle Elum-Roslyn School District. To date we have captured and marked with 

GPS collars 4 adult and 2 subadult male and 4 adult female cougars. GPS transmitter collars 

collect GPS coordinates at 4-hour intervals throughout the year. This data is plotted onto GIS to 

assess proximity to human residence, planned development, recreational centers, and to assess 

predation events and habitat use patterns. This investigation is used to engage students in an 

experiential learning activity whose focus is application of technology and learning about their 

ecological and social community. Students in kindergarten to senior high help collect and 

analyze data. Junior-Senior students in Advanced Placement Biology assist with cougar capture 

and marking efforts and correlate location data with GIS habitat, topographic, and human 

residence parameters. They will use DNA isolated from cougar scats to determine species and 

gender of animals depositing scats while 8 th grade students analyze scats for contents to correlate 

food habits with gender of cougars. Elementary students learn plant identification for plotting 

habitat types and learn animal track identification for reporting locations of carnivores and 

ungulate prey species near their residence. Students count ungulates along bus routes for longterm 

monitoring of prey distribution in relation to seasons and development. Students and 

community member conduct tests of GPS collars to assess influences of vegetative and 

physiographic conditions on satellite acquisition rate and accuracy. Students are assessed on 

their abilities to collect qualitative and quantitative data. Community members help collect data 

and help train students in outdoor and data collection skills. Central Washington University 

incorporates Project CAT objectives into training teachers. Information on ungulate habitat and 

cougar travel corridors is shared with community planners to incorporate into planning processes 

to minimize human-cougar interactions. 


MONITORING MOUNTAIN LIONS IN THE TUCSON MOUNTAIN DISTRICT OF 

SAGUARO NATIONAL PARK, ARIZONA, USING NONINVASIVE TECHNIQUES 

LISA HAYNES, Wild Cat and Carnivore Studies, 133 W. 2 nd St., Tucson, AZ 85705, USA, 

email: lynxrufus@earthlink.net 

DON SWANN, Saguaro National Park, 3693 S. Old Spanish Trail, Tucson, AZ 85730, USA, 

email: Don_Swann@nps.gov 

MELANIE CULVER, Arizona Cooperative Fish and Wildlife Research Unit, 104 Biosciences 

East, University of Arizona, Tucson, AZ 85721, USA, email: culver@ag.arizona.edu 

Abstract: This presentation will summarize a three-year effort to noninvasively monitor 

mountain lions in the Tucson Mountain District of Saguaro National Park, Arizona from 2001 

through the spring of 2003. Park managers are concerned about this mountain lion population, 

because the Tucson Mountains are becoming surrounded by human development. The continued 

existence of the population is threatened due to habitat loss, potential inbreeding, and disrupted 

demographics. Noninvasive methods used to monitor mountain lions included track surveys, 

infrared-triggered cameras, and molecular genetic analysis of hair (collected from hair snares) 

and scat (feces). In the first two years, 2001 and 2002, we documented a total of 19 sets of 

mountain lion tracks found during 2 winter surveys of 30 transects and during seasonal surveys 

of 4 transects. From track data we determined that mountain lions are consistently detected in 

most areas of the park; however, preliminary evidence suggests a paucity of adult males. One of 

the most important aspects of this project is the opportunity for educating volunteers from the 

general public who participate in the track surveys. In 2002, 31 of 35 hair snares distributed 

throughout the park had hair deposited on them. Planned genetic analysis will determine 

efficacy of using hair snares and scat to obtain biological information on mountain lions. We 

will present additional data from the 2003 track survey and we will provide recommendations for 

future monitoring and research efforts. 


141

142 

ESTIMATING COUGAR ABUNDANCE USING PROBABILITY SAMPLING: AN 

EVALUATION OF TRANSECT VERSUS BLOCK DESIGN 

CHUCK R. ANDERSON, JR., Wyoming Cooperative Fish and Wildlife Research Unit, Box 

3166, University Station, Laramie, WY 82071. USA, email: cander@uwyo.edu 


University Station, Laramie, WY 82071, USA, email: flindzey@uwyo.edu 

NATE P. NIBBELINK. Wyoming Geographic Information Science Center, University of 

Wyoming, Box 4008, University Station, Laramie, WY 82071, USA, email: 

nathan@uwyo.edu 

Abstract: We used GPS data records of cougar track sets (n = 5-6 locations/night) to evaluate 

accuracy and precision of Transect Probability Sampling (TPS) and Block Probability Sampling 

(BPS) from spatial simulations of varying cougar densities, sampling efforts, and number of 

track set nights. GPS data records yielded 446 1-night track sets and 225 2-night track sets from 

12 cougars (2 adult males, 6 adult females, and 4 subadults) for simulations. Accuracy and 

precision of TPS and BPS estimates generally improved with increased cougar density, sampling 

effort, and number of track set nights, but TPS estimates were vulnerable to extremely short 

track sets (e.g., cougars at kill sites) and BPS estimates were exceedingly imprecise. To address 

these problems, we adjusted TPS estimates based on the proportion of cougar track sets 

estimated to be at kill sites and used bootstrap techniques to estimate 90% confidence intervals 

(CIs) around BPS estimates. TPS estimates adjusted for cougars at kill sites typically improved 

accuracy, precision, and estimator reliability (CIs approaching 90% coverage). Bootstrapping 

greatly reduced variance around BPS estimates but exaggerated precision (i.e., CI coverage 

typically below 90%), likely due to low cougar detection rates. Comparisons of adjusted TPS 

and BPS estimates suggested higher cougar detection rates and improved accuracy from TPS 

surveys, with more reliable CI coverage. TPS simulations suggested reliable cougar population 

estimates could be obtained from high-effort surveys (~2 km transect spacing) regardless of 

cougar density or number of track set nights, or from medium-effort surveys (~3 km transect 

spacing) of medium-high density populations (2.3-3.5 independent cougars/100 km 2 ) sampling 

2-night track sets. Ninety-percent CIs suggested population changes of 27-30% could be 

detected using high effort surveys of 1-night track sets, 20-24% from medium effort surveys of 

2-night track sets, and 15-18% from high effort surveys of 2-night track sets. Because of the 

time and expense required to conduct high effort TPS surveys, we propose sampling cougar track 

sets without intense tracking efforts and applying perpendicular track lengths we measured to 

estimate cougar population parameters. 


EVALUATING MOUNTAIN LION MONITORING TECHNIQUES IN THE GARNET 

MOUNTAINS OF WEST CENTRAL MONTANA 

RICH DeSIMONE, Montana Fish, Wildlife & Parks, 1420 East Sixth Avenue, Helena, MT 

59620, USA, email: rdesimone@state.mt.us 

Abstract: Research began in 1998 to document characteristics of a hunted mountain lion 

population and develop survey techniques to detect trends in lion abundance. Efforts to capture 

and radio-collar mountain lions have focused on the 850 km 2 eastern half of the Garnet 

Mountains where lion hunting was suspended from 2000 to 2002, allowing the lion population to 

increase. Lion hunting will resume, reducing the number of lions in the study area. Fluctuations 

in a known lion population will provide the opportunity to determine the sensitivity of population 

indicators to changes in lion abundance. Mountain lion population trend indicators being 

evaluated include lion track survey-routes and statewide telephone surveys of houndsmen and 

deer hunters. Eleven lion track snow-survey-routes totaling approximately 105 km were 

established in 2000 throughout the study area to determine the relationship between lion track 

density and the actual density of lions. Track densities ranged from 0 to 2 per 10 km. 

Preliminary results indicate that the densities of lion tracks recorded in different portions of the 

study area correlate with the densities of lion home ranges. A statewide telephone survey of 

houndsmen began in 2001 with approximately 300 houndsmen interviewed annually. 

Houndsmen took fewer days of hunting to tree lions (3 days) and encounter a lion family group 

(8 days) in northwest Montana, while in eastern Montana houndsmen took 10-35 days to tree a 

lion and 35-45 days to encounter a family group. Starting in 2001, the statewide telephone 

survey of deer hunters included asking hunters if they observed lions. The percentage of deer 

hunters observing lions ranged from 4% in northwest Montana to less than 1% in eastern 

Montana. Seventy lions have been captured and radio-collared. Eleven of 26 radioed kittens died 

during their first year of life. Malnutrition due to orphaning was the most common cause of 

death. Hunters harvested adult and subadult lions at a high rate. In portions of the study area 

where hunting was allowed, hunters harvested an average of 63% of the radioed lions annually 

from 1998 to 2001. Overall, 36 of 38 radioed lion deaths were human related. 


143

144 

PRESENCE AND MOVEMENTS OF LACTATING AND MATERNAL FEMALE 

COUGARS: IMPLICATIONS FOR STATE HUNTING REGULATIONS 

TONI K. RUTH, Wildlife Conservation Society, 2023 Stadium Dr. Suite 1A, Bozeman, MT 

59030, USA, email: truth@montanadsl.net 

KERRY M. MURPHY, Yellowstone Center for Resources, P.O. Box 168, Yellowstone National 

Park, Mammoth, WY 89210, USA, email: kerry_murphy@nps.gov 

POLLY C. BUOTTE, Wildlife Conservation Society, 2023 Stadium Dr. Suite 1A, Bozeman, MT 

59030, USA, email: polly_thornton@hotmail.com 

Abstract: Established in the early 1970’s, the regulation of cougar harvest through hunting 

seasons and quotas contributed to increases in cougar abundance and distribution in most 

western states during the past 30 years. Today, 10 of 12 western U.S. states regulate cougar 

harvest through hunting seasons and quotas, which vary by state, year, and in season length and 

quota numbers. California prohibits cougar hunting and Texas allows unlimited hunting and 

therefore lacks regulations. Three of 11 states that allow cougar hunting regulate the harvest of 

female lions through subquotas. Two states that do not regulate female harvest do not allow the 

use of hounds for hunting cougars. Hunting regulations in 9 of 11 states prohibit killing spotted 

kittens and females with spotted kittens. Only one state, Montana, requires hunters to backtrack 

lactating females that have been killed in order to locate dependent young. While regulations 

prohibiting the take of maternal females and affording protection to nonmaternal females through 

subquotas should remain in place, little information has been provided to hunters, guides and 

outfitters, or state managers on the proportion and movements of lactating and maternal females 

which may be encountered during hunting seasons. We examined reproductive data for a 

moderately hunted cougar population (1988-1992) and a primarily non-hunted population (1998- 

2002) on the Northern Range of Yellowstone National Park. Proportion of females with 

dependent (pre-dispersal) offspring was calculated across winters. We summarized breeding, 

denning, and lactation chronology for maternal females from both study periods. Peak breeding 

occurred in March through May and denning followed approximately 3 months later, peaking in 

June through August. Given a 4-month lactation period, proportion of lactating females (n = 19 

known date births or births estimated to

MYSTERY, MYTH AND LEGEND: THE POLITICS OF COUGAR MANAGEMENT IN 

THE NEW MILLENNIUM 

RICK A. HOPKINS, Live Oak Associates, Inc., 6830 Via Del Oro, Suite 805, San Jose, CA 

95119, USA, email: rhopkins@loainc.com 

Abstract: The cougar as America’s cat is a large, ghost like predator that usually hunts game as 

large as or larger than it is. As Teddy Roosevelt noted in the late 1800’s, “No American beast 

has been the subject of so much loose writing or of such wild fables as the cougar”. More than 

100 years and dozens of scientific studies later, we are no better off in unraveling the “loose 

writing” and in some cases, the management objectives for this predator, than we were at the 

beginning of the twenty century. The life style of the cat has long resulted in polarized attitudes 

toward the development of policies for its management throughout North America. The general 

“truth” that has evolved with wildlife managers regarding increasing cougar numbers throughout 

the West over the last 2 to 3 decades is believed to be born from 30 years of research. While 

numbers of cougars may have increased in portions of their range over the last 2-3 decades, the 

general perception that cougars are more abundant in the western U.S. is based not on empirical 

data, but one based more on oral traditions passed on from one wildlife professional to another. 

Cougar management in the last two decades focused on the impact of prey populations and 

depredation of livestock more than on direct encounters with humans. However, an increase in 

human attacks in the 1990’s has not only heightened public awareness of cougars, but appears to 

have explicitly shifted management in some western states to focus more on “controlling” the 

species. Lost in this debate and the objectives of the management of the species is the 

preservation of those elements that will truly lead to its conservation. California represents an 

interesting living lab, as cougars have not been sport hunted in this state since 1972; it also 

supports one of the largest cougar populations in the U.S. if not the largest, and clearly supports 

the largest human population. The lessons learned in California can serve as a model for 

continued efforts to focus management objectives on the conservation of the species and not 

solely on equating management with harvest as is so often done. 


145

146 

RECONCILING SCIENCE AND POLITICS IN PUMA MANAGEMENT IN THE 

WEST: NEW MEXICO AS A TEMPLATE 

KENNETH A. LOGAN, Carnivore Researcher, Colorado Division of Wildlife, 2300 South 

Townsend Avenue, Montrose, CO 81401, USA, email: Ken.Logan@state.co.us 

LINDA L. SWEANOR, Scientist, Wildlife Health Center, University of California, TB128, Old 

Davis Road, Davis, CA 95616, USA, email: lsweanor@mindspring.com 

MAURICE G. HORNOCKER, Senior Scientist, Wildlife Conservation Society, Box 929, 

Bellevue, ID 83313, USA 

Abstract: The puma is the only large obligate carnivore thriving today in self-sustaining populations 

distributed across western North America. As such, the puma contributes to ecosystem integrity because 

the puma: 1) strongly influences energy flow and nutrient cycling; 2) is a strong natural selective force on 

prey animals; 3) modulates prey population dynamics; 4) indirectly affects herbivory on plant 

communities; 5) indirectly influences competition among herbivores; and 6) competes with other 

carnivores. Furthermore, because persisting puma populations depend on expansive, connected wild 

landscapes with thriving prey populations, the puma is also a potential focal species for designing nature 

reserve networks. Wildlife managers have the responsibility of weighing the natural value of the puma 

with the diverse needs of people. Yet, their tools for scientific puma management are crude mainly 

because pumas are very cryptic and exist in very low population densities. People in New Mexico 

identified 10 puma management issues: 1) pumas kill livestock and threaten rancher’s livelihoods; 2) 

pumas kill deer that could be taken by hunters; 3) pumas threaten conservation of endangered populations 

of mountain sheep; 4) some pumas threaten public safety; 5) sustainable puma hunting is desirable; 6) 

puma hunting should focus on taking males and protecting females and cubs; 7) hunting pumas with dogs 

is undesirable; 8) puma hunting is undesirable; 9) increased human development threatens puma 

conservation; 10) diverse interests make puma management difficult. Unknowns and uncertainties 

specific to puma management included: 1) number of pumas in populations; 2) population trends; 3) 

population growth rates; 4) population responses to management prescriptions; 5) effects of hunter 

selection; 6) density distributions; 7) age and sex structure of populations; 8) reproductive rates; 9) agespecific 

survival rates; 10) immigration and emigration rates; 11) validity of puma population simulation 

models. These unknowns and uncertainties along with the broad diversity of human values toward the 

puma make management very difficult and challenge the professional integrity of agencies. In New 

Mexico, we developed a robust, biologically sound, adaptive puma management structure that considers 

the role of the puma in ecosystems, the needs of people, and the unknowns and uncertainties in puma 

management. We called this structure Zone Management. Zone Management uses zones with lethal 

control, sport-hunting, and refuges. Control zones allow experimental puma control in focal areas to 

protect private property, human safety, endangered species, or game animals. Hunting zones allow sporthunting 

opportunity sustained by quotas on the number of pumas that can be killed, with emphasis on 

protecting females and cubs. Refuge zones (i.e., no hunting zones) are >3,000 sq. km and act as biological 

savings accounts that assist wildlife managers by countering mistakes made in the control and hunt zones, 

allowing natural selection to occur in puma populations, and providing numeric and genetic augmentation 

of human impacted zones via puma dispersal from refuges and immigration into human exploited zones. 

The zone management structure uses the source-sink metapopulation paradigm we developed for pumas 

in New Mexico. 


COMMUNITY-BASED CONSERVATION OF MOUNTAIN LIONS 





Abstract: The western United States is experiencing a rapid growth in human population, with a 

commensurate loss and fragmentation of wildlife habitat. As the only top predator with viable 

populations throughout the West, the long-term conservation of this species is vital for 

maintaining the health and integrity of the region’s native ecosystems. Unfortunately, 

populations of mountain lions in areas subjected to intensive human development or activity may 

become threatened unless conservation efforts are implemented and realized. For example, in 

several regions of California, including the West slope of California’s Sierra Nevada Mountains, 

the number of mountain lions killed as the result of conflicts with domestic animals (e.g., goats 

and pets) has increased dramatically over the past decade. According to California Dept. of Fish 

and Game, mountain lions may be extirpated from this area within 40 years due to habitat loss 

and excessive human caused mortality. Accordingly, a substantial policy shift is needed. In other 

areas, such as Southern California, few lions remain in fragmented wildlands, and different 

conservation strategies are required to reach urban and suburban residents who live on the edge 

of wildlife areas. We review efforts to conserve mountain lions at the community level and 

elaborate new approaches that stress science to establish a factual basis for dialogue, community 

involvement to identify shared goals, and developing partnerships with diverse organizations and 

professions to broaden conservation efforts. 


147

148 

PUMA MANAGEMENT IN WESTERN NORTH AMERICA: A 100-YEAR 

RETROSPECTIVE 

STEVEN TORRES, California Department of Fish and Game, 1416 Ninth Street, Room 1342A, 

Sacramento, CA 95814, USA, email: storres@dfg.ca.gov 

HEATHER KEOUGH, Utah State University, Logan, UT 84322, USA 

DEANNA DAWN, South Dakota State University, Brookings, SD 57007, USA 

Abstract: Puma (Puma concolor) populations have had a diverse and long history of 

management in western North America. For the most part of the last century, pumas were a 

bountied predator. By the early 1970s, they had transitioned to game mammal status. In the 

period since bounties ended, most states and provinces have reported increased puma activity 

that has been simultaneous with increased human populations and land conversion. We will 

present an analysis of the political and biological effects influencing puma populations during 

this period to provide perspective on the potential effects of bounty removals as they may relate 

to hypothesized increased populations in the latter part of the last century. This presentation will 

also explore the changing philosophy of predator management and the importance of maintaining 

predator-prey systems and redefining puma management to include their beneficial role in 

defining large blocks of habitat and movement corridors. 


USING COUGARS TO DESIGN A WILDERNESS NETWORK IN CALIFORNIA’S 

SOUTH COAST ECOREGION 

PAUL BEIER, Northern Arizona University, Flagstaff, AZ 86011, USA, email: 

Paul.Beier@nau.edu 

KRISTEEN PENROD, South Coast Wildlands Project, PO Box 2493, Monrovia, CA 91016, 

USA, email: Kristeen@scwildlands.org 

Abstract: The groundbreaking “Missing Linkages” report published in fall 2001 

(www.scwildlands.org) identified over 200 linkages needed to prevent isolation of wildlands in 

California. South Coast Wildlands Project (SCWP) immediately spearheaded an effort to 

prioritize, protect and (where necessary) restore linkages in the South Coast Ecoregion. SCWP 

first assessed the ecoregion’s 69 linkages with respect to biological irreplaceability (size and 

quality of core areas served by a linkage were important criteria) and vulnerability (to 

urbanization and roads). This process identified 15 linkages as top priorities. We are now in the 

process of conducting a series of action workshops for each linkage. At the first workshop, local 

biologists, government agencies and conservation NGO representatives developed lists of focal 

species and ecological processes that a linkage is intended to serve. Thus, although carnivores 

helped to initially identify important linkage areas, we are designing each linkage to serve 

broader biodiversity goals. Our personnel are researching the needs of the focal species, 

obtaining high-resolution photographs and parcel maps, and conducting field visits. One or more 

linkage designs will be presented at a second workshop, where participants will volunteer for 

various tasks (e.g., procuring easements, acquiring land, changing zoning, restoring habitat, or 

mitigating transportation projects) to preserve and enhance the linkage. By partnering with 

agencies and NGOs from the start rather than developing a plan on our own and asking others to 

unite under us, our effort has attracted funding and cooperation from diverse sources and is 

making rapid progress. 


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150 

MOUNTAIN LIONS AND BIGHORN SHEEP: FACING THE CHALLENGES 



JOHN D. WEHAUSEN, U.C. White Mountain Research Station, 3000 E. Line Street, Bishop, 

CA 93514, USA 

Abstract: Mountain lions (Puma concolor) and bighorn sheep (Ovis canadensis) have coevolved 

as predator and prey. Bighorn sheep populations have declined over the past several 

centuries across their range due to a variety of anthropogenic causes, and have been the subject 

of extensive translocation efforts in recent years in an effort to reestablish populations in historic 

habitat. In the past several decades mountain lion predation has also been implicated in the 

decline of several populations of endangered bighorn sheep, including the federally listed 

Peninsular (O. c. cremnobates) and Sierra Nevada populations in California and state listed 

desert bighorn sheep (O.c. mexicana) populations in New Mexico. We hypothesize that this 

recent phenomenon has resulted from land and wildlife management practices that have affected 

both species. Restoring the natural relationship between mountain lions and bighorn sheep 

presents both biological and ethical challenges. We discuss the current status of management 

efforts, review several hypothesis of why predation has become a limiting factor for bighorn 

sheep recovery, and discuss current and potential management options. 


FACTORS AFFECTING DISPERSAL IN YOUNG MALE PUMAS 

JOHN W. LAUNDRÉ, Instituto de Ecologia, A.C, Km 5 a Carr. de Mazatlán s/n, C.P 34100, 

Durango, Dgo México, email: launjohn@prodigy.net.mx 

LUCINA HERNÁNDEZ, Instituto de Ecologia, A.C, Km 5 a Carr. de Mazatlán s/n, C.P 34100, 

Durango, Dgo México, email: lucina@fauna.edu.mx 

Abstract: Numerous studies have demonstrated that nearly all young male pumas disperse from their natal home 

range area while most females are philopatric. There are 2 hypotheses for the driving force behind dispersal in 

young male pumas. The first is the competition (aggression avoidance) model where dispersal is because of 

competition between young males and their fathers/incoming transient males for mates and resources. The second is 

the inbreeding model where young males are thought to disperse to avoid inbreeding with their mothers/sisters. 

Under each model, specific predictions can be made to test their validity. Under the competition model, we predict 

that there should be physical conflicts between sons and their fathers, including infanticide and that competitive 

ability should increase with age and thus, the longer time and further distance they are from their home area. Thus, 

dispersal distances should reflect their competitive ability, i.e. few males will establish territories close to their natal 

home range. Under the inbreeding avoidance model, we predict fewer physical conflicts (young males leave on 

their own), no inbreeding between fathers and daughters, which is genetically equivalent to sons mating with their 

mothers, and no males should establish their territories adjacent (1-2 home range diameters) to their natal home 

range. We tested these predictions with dispersal data from our study in southern Idaho/northwestern Utah and 

published data. We refuted the inbreeding model because resident males do fight and kill their male offspring, 

resident males do mate with their daughters, and there is a high percent of males that establish their territories within 

2-4 HRDs of their natal home ranges. Our data supported the competition model with an increase in frequency of 

dispersal distances at >2 diameters. We conclude that young males are forced out of their natal home range by their 

fathers or incoming males who, by default, will be older and stronger. We propose that they continue to disperse 

until they gain enough weight and experience to successfully takeover a territory. 

Key Words: male pumas, dispersal, inbreeding, competition, Idaho, Utah 

Numerous studies have demonstrated 

that nearly all young male pumas disperse 

from their home range area while most 

females are philopatric (Ross and 

Jalkotzy1992, Sweanor et al. 2000). There 

are 2 hypotheses for the driving force behind 

dispersal in young male pumas that have 

been developed considering the various 

costs and benefits to dispersing vs 

philopatric behaviors (Shields 1987). The 

first is the competition model where 

dispersal is motivated by interference 

competition and aggression between young 

males and their fathers/incoming transient 

males for mates and resources (Dobson 

1982, Moore and Ali 1984). The second is 

the inbreeding model where young males are 

thought to disperse to avoid inbreeding 

151 


primarily with their mothers/sisters (Wolfe 

1994). There have been extensive reviews 

and criticisms of both hypotheses and it 

continues to be a hotly debated topic (See 

Shields 1987 for a review). Relative to 

pumas, discussions of dispersal have 

primarily centered around dispersal ages and 

distances but little on the driving force 

behind dispersal, especially for young 

males. Sweanor (1990) reported on 

dispersal in male pumas in New Mexico and 

concluded that the likely driving force was 

competition among males. However, later, 

Logan and Sweanor (2001) concluded that 

male pumas were dispersing primarily to 

avoid inbreeding. Thus at this time it is 

uncertain which hypothesis might best 

explain dispersal in young male pumas.

152 DISPERSAL IN MALE PUMAS · Laundré and Hernández 

Under each model, specific predictions 

can be made to test their application to male 

pumas. Under the competition model, we 

predict that there should be physical 

conflicts between adult males/kittens 

(infanticide), adult males/juveniles, 

including father/son, and adult male/adult 

male. Older, more experience males should 

win these conflicts and competitive ability 

of dispersers should increase with age and 

thus, the longer time and further distance 

they are from their home area (Sweanor 

1990). Thus, another prediction for this 

model is that dispersal distances should 

reflect their competitive ability, i.e. few 

males will establish territories close to their 

natal home range. Under the inbreeding 

avoidance model, we predict less physical 

conflicts (young males leave on their own), 

no inbreeding between fathers and 

daughters, which is genetically equivalent to 

sons mating with their mothers, and no 

males establishing territories within a 

minimum of 2 home range diameters of their 

natal home range (Sweanor 1990). We 

attempted to test these predictions with data 

from our study in southern 

Idaho/northwestern Utah and published data 

from a variety of other studies of pumas. 

We recognize that it is often difficult to 

obtain good dispersal data on a species such 

as the puma but combining the data from the 

various studies that exist should provide us 

the best opportunity to examine possible 

causes of dispersal in male pumas. 

METHODS 

To test the predictions made, we used a 

combination of data from our long-term 

study of pumas and published data on other 

puma studies. In some of these studies, 

pumas were protected from hunting while in 

the others they were exposed to sport 

harvest. However, in the case of the 

protected populations, the protection 

extended only to the limits of the study 

areas, which ranged in size from ≈ 500 to 


2,000 km 2 or a maximum diameter of 50 

km. Because in all the studies, most young 

male pumas dispersed to outside of the 

designated study areas, we consider that 

pumas were dispersing under unprotected 

conditions. Consequently, we did not 

subdivide the studies into protected and 

unprotected populations. 

Our study of a harvested puma 

population was conducted in southeastern 

Idaho and northwestern Utah. The study 

area was approximately 2,000 km 2 and 

consisted of 5 small mountain ranges 

(approximately 1,000 km 2 total area) 

separated by valleys where human activity 

predominated. Over 16 years, we conducted 

intensive capture efforts each winter and 

with the help of trained dogs, we captured 

and marked approximately 150 pumas. We 

documented male-male conflicts based upon 

our intensive field efforts. We obtained 

dispersal data of young radio collared male 

pumas primarily from hunter returns. We 

measured straight-line dispersal distance 

from the center of the natal home range to 

the point of harvest. In addition to our data, 

we searched the published literature for 

accounts of male-male conflicts, incidences 

of inbreeding, and estimates of dispersal 

distances. For male conflicts, we considered 

3 levels of conflict, adult male/kitten 

(infanticide), adult male/subadult male, and 

adult male/adult male. Incidences of 

inbreeding were obtained via field 

observations and, in some cases, genetic 

testing. 

For dispersal, we expressed male 

dispersal distances from natal home ranges 

in multiples of the average male home range 

diameter (HRD) (Waser 1985). We 

calculated HRDs for each study assuming a 

circular home range (Sweanor et al. 2000). 

We then estimated the cross study HRD 

average and then divided dispersal distances 

of each study by the cross study mean HRD. 

Although we assumed the mean of the

different HRD estimates would be a better 

estimate of the true HRD for male pumas 

overall, we recognize that the variation 

among studies might represent true regional 

differences and not just random variation 

around the population mean. Thus, we also 

expressed dispersal distances of each study 

in multiples of the individual study estimates 

of HRD. 

We plotted the frequency distributions of 

dispersal distances expressed as multiples of 

HRDs for both individual study estimates 

and for the across study estimate of HRDs. 

To test if competition drives male dispersal, 

we compared the observed percent 

frequency distribution of dispersal distances 

(x) with that expected (f(x)) calculated from 

Equation 1 (Miller and Carroll 1989). 

Equation 1: 

x 

F(x) = p(x) ∏ [1-p(x-i)] 

i = 1 

In this equation, f(x) is the probability an 

animal will establish a territory at x 

dispersal distance units, in this case HRDs, 

from the natal home area. The value p(x) is 

the probability that an animal will obtain a 

territory at x dispersal distances and varies 

from p(0), the probability the animal will not 

leave its natal area, to a maximum value, 

usually represented by the average mortality 

rate (Waser 1985). The quantity, 1-p(x-i), is 

the probability of continuing through 

previous habitats. We selected the model of 

Miller and Carroll (1989) over other 

geometric models such as that of Waser 

(1985 and 1987) because Equation 1 

incorporates changes over distance in the 

probability of an animal securing a territory. 

We considered the model of Miller and 

Carrol (1989) more realistic because based 

on all the reported data (Logan and Sweanor 

2001), the probability of a young male 

DISPERSAL IN MALE PUMAS · Laundré and Hernández 153 


setting up his territory within his natal area 

(p(0)) is zero. 

We estimated 2 expected frequency 

distributions calculated from Eq. 1. The 

first was an estimation of what we would 

expect under the competition model. We 

assumed that once young males start 

dispersing, their competitive ability should 

increase with more time, and thus, distance 

they disperse until they are equally 

competitive for any open territories (Logan 

and Sweanor 2001). Logan and Sweanor 

(2001) reported that dispersing males in 

their study rarely move constantly but move 

from one transient home range to another. 

However, the amount of time and distance 

they traveled between the natal and 

independent home range varied extensively. 

Of 4 dispersing males Logan and Sweanor 

(2001) monitored, the dispersal time 

averaged 211 + 192 days and straight-line 

dispersal distances averaged 143.2 + 26.5 

km. Based on these estimates, male pumas 

will move on average 0.7 km per day. They 

disperse at approximately 16-18 months 

(Anderson et al. 1992, Ross and Jalkotzy 

1992, Hemker et al. 1984, Beier 1995, 

Maehr et al. 1991, Logan and Sweanor 

2001) and in 5 months would move 

approximately 100 km which, based on 

results presented later, represents 

approximately 5 HRDs. At 21-24 months 

they are near their full body mass (Laundré 

and Hernández 2002) and we assumed to be 

competitive for territories. Thus the 

competition model incorporated a linear 

change in p(x) from p(0) = 0 to the average 

mortality rate over the first 5 HRDs (x = 1 

to 5) and then after that, it was held constant 

at the average mortality rate. 

For the inbreeding model, we set p(x) = 

0.0 for the first few HRDs (depending on the 

average dispersal distance of females) and 

after that, set it to the average mortality rate; 

young male pumas should move beyond the 

areas where there is a high probability of


mating with sisters or mothers that have 

shifted their home range areas. This 

minimal distance should be represented by 

the average dispersal distance of females 

(sisters) from their natal home range. 

RESULTS 

Prediction #1: Male conflicts 

Relative to levels of adult male/kitten 

conflict or infanticide, we found ample 

evidence for infanticide in our study and in 

the literature. We recorded 4 cases of adult 

male pumas killing kittens (9 kittens total) in 

our study area. Robinette et al. (1961) cited 

reports from Utah of young pumas being 

killed by adult males on 2 occasions. 

Hornocker (1970) reported 1 instance of an 

adult male killing 2 kittens in his study area 

in central Idaho. Lindzey et al. (1989) 

reported 4 kittens being killed by pumas in 

southern Utah. However, they were not sure 

of the sex of the responsible animals. 

Spreadbury et al. (1996) documented 2 male 

kittens being killed by a transient male in 

British Columbia. Logan and Sweanor 

(2001) reported 12 kittens being killed by 

adult males in their study in New Mexico. 

With respect to adult male/subadult male 

conflicts, Lindzey et al. (1989) reported a 

transient male being cannibalized by a 

resident male. Murphy (1998) reported 2 

incidences of territorial males killing 

yearling males in his study in Yellowstone 

National Park. Logan and Sweanor (2001) 

reported 4 mortalities of younger (14.3-21 

months) males being killed by older males, 

including 1 instance of a father killing his 

son. 

Although Hornocker (1970) and 

Seidensticker et al. (1973) did not report 

conflicts among adult male pumas, various 

other investigators reported it as rather 

common. In our area, adult males we 

captured often had facial scars and torn ears, 

presumably from conflicts with other adult 

males or, perhaps, females. Sitton and 


Weaver (1977) reported 2 adult pumas being 

killed by other adult pumas as a result of 

fighting. However, the sex of the 

combatants was unknown. McBride (1976) 

reported 2 incidences of adult males with 

extensive facial injuries and 1 instance of an 

adult male being killed by another. Murphy 

(1998) reported an adult immigrant male 

being killed by the resident male. Logan 

and Sweanor (2001) found 5 adult males 

killed by other males. They also reported 

that 4 of the 5 winners outweighed the 

losers, 2 of the 5 losers were killed by older 

animals, and 2 of the other 3 were 8 and 12 

years old, well past their prime. 

Prediction # 2: occurrence of inbreeding 

Concerning the presence of inbreeding 

within puma populations, it is well known 

that the Florida panther is highly inbred, 

including first order matings (Roelke et al. 

1993, Barone et al. 1994 and others). This 

indicates that there is no innate “aversion” to 

inbreeding. Logan and Sweanor (2001) 

reported an average residency of males of 6 

years (maximum = 8 yrs) in their study in 

New Mexico. If we assume an average of 2 

years per litter (Logan and Sweanor 2001), 

males were resident on average for at least 3 

generations. Given that females are 

primarily philopatric (Logan and Sweanor 

2001), and there is no aversion to inbreeding 

with their fathers, we predict that with this 

length of residency a father would have the 

opportunity to mate twice with his daughters 

and once with granddaughters. If there is an 

innate aversion to inbreeding, such matings 

should not occur. Logan and Sweanor 

(2001) report that 2 females mated with their 

fathers, 1 of them mated with her father 

twice, supporting the prediction based on no 

aversion to inbreeding. 

Prediction # 3: Dispersal distances 

For dispersal distances of males, we 

limited our analysis to animals that were 

known to have set up territories or were


Table 1. Average dispersal distances (km), home range sizes (HRS; km 2 ), and home range 

diameters (HRD; km) of male pumas from various studies. The home range sizes are based on 

resident males in the study areas. Dispersal distances are only of individuals who were known to 

establish territories after dispersing or were at least > 24 months old before being killed. 

Source n Distance H.R. Size HRD 

Lindzey et al. (1989 & 1994) 4 142.2 731.0 30.5 

Logan et al. (1986) 3 175.7 320.0 20.2 

Spreadbury et al. (1996) 4 74.5 132.3 13.0 

Hornocker (1960) & 

Seidensticker et al. (1973) 

5 102.4 379.5 22.0 

Ashman et al. (1983) 6 48.9 355.2 21.3 

Logan and Sweanor (2001) 13 128.2 187.1 15.4 

This study 13 190.6 302.5 19.6 

Averages 123.2 343.9 20.3 

at least >24 months old at the time they were 

killed (Sweanor et al. 2000). We found 

dispersal distances for 48 male pumas from 

7 different studies, including 13 estimates 

from our study area (Table 1). We did not 

use the dispersal distances reported by Beier 

(1995) because his study area, although 

large (2070 km 2 ), was completely 

surrounded by urban development, which 

limited dispersal opportunities. Dispersal 

distances ranged from 31 km (Spreadbury et 

al. 1996) to a maximum of 550 km (our 

study). Estimates of resident home range 

sizes of males in these studies varied 

between 132.3 to 731 km 2 (Table 1). The 

mean of these averages was 343.9 km 2 with 

a HRD of 20.3 km. Dispersal distances 

varied from 0.9 to 26 HRDs with a mean of 

6.0 HRDs (Fig. 1). The majority (≈ 50 %) 

of dispersal distances were between 2 and 4 

HRDs with a second peak (25 %) between 

8-9 HRD’s. There was a noticeable 

reduction of dispersal settlements between 5 

and 7 HRDs (Fig. 1). 

Based on the individual estimates of 

home range size, the frequency of HRDs 

changed in some categories (Fig. 1). 

However, the pattern was similar to the 

distribution based on the overall average 

HRD, e.g. the majority of the dispersal 


distances were between 2 and 4 HRDs. The 

one difference was the appearance of more 

settlements in the 5-7 HRD range (Fig. 1) 

We found estimates of male mortality 

rates from 4 other studies (Ashman et al. 

1983, Ross and Jalkotzy 1992, Anderson et 

al. 1992, Logan and Sweanor 2001) plus our 

Figure 1. Frequency distribution of 

dispersal distances, expressed as multiples 

of the cross study average home range 

diameter (HRD) and as multiples of the 

individual study HRDs. The two curves 

represent the expected frequency 

distribution for the competition and the 

inbreeding models.


estimate and they ranged from 9.0 % to 31.0 

% with an average mortality rate of 20.5%. 

Based on movement distances and times 

presented in Logan and Sweanor (2001), we 

assumed the competitive ability of a young 

male puma would be at its maximum by 5 

HRDs. Thus, in Eq. 1, for the competition 

model, we let p(x) vary from 0.04 to 0.18 

over the first 4 HRDs (x = 1 to 4) and then 

after that, let it be a constant at 0.205, the 

average mortality rate. The resulting curve 

underestimated the number of settlements in 

the second and third HRD but the pattern fit 

the data relatively well in the first 4 HRDs. 

In contrast it predicted more settlements in 

the 5-7 HRD range and less in the 8-9 range 

than what we found (Fig. 1). 

For the inbreeding model, we found in 

our study and in that of Logan and Sweanor 

(2001), approximately 42.0 % of the young 

females dispersed an average of 2 HRDs 

(range = 1 to 8 HRDs, n = 14). Based on 

this, a dispersing male would almost be as 

likely to mate with a sister within the first 2 

HRDs as in his natal home range. Also, 

considering that resident females can easily 

shift their home range area by at least 1 

HRD during their reproductive years, we 

concluded that to avoid inbreeding, a male 

should disperse at least a minimum of 3 

HRDs before seeking a territory. 

Consequently, we set p(x) = 0.0 for x = 1 to 

2 and after that, set it to the average 

mortality rate of 20.5 %. As 25.0 % of the 

dispersing males settled within the first 2 

HRDs (Fig. 1), this model initially did not fit 

the data. It also predicted a higher 

occurrence of settlements in the third and 

fourth HRDs than what we found. From the 

fifth HRD this model patterned after the 

competition model. 

DISCUSSION 

Regarding prediction # 1, we found 

ample evidence there are high levels of 

conflict between adult territorial males and 

all age classes of non-territorial males 


(kittens to adults, including between fathers 

and sons). These conflicts at times appear to 

be over food resources but primarily are 

over breeding resources. Also in almost all 

cases, the older, heavier, more experienced 

male wins the conflict. These data support 

the competition hypothesis for dispersal 

because if young males do not disperse they 

will be killed by their fathers or nearby 

resident males, all of whom are bigger and 

more experienced. 

For prediction # 2, although it can be 

argued that the Florida pumas inbreed 

because it is a closed population, it still 

refutes the myth of a natural aversion to 

inbreeding. The innate aversion to close 

inbreeding is the driving force behind the 

inbreeding avoidance model and, for the 

model to work, should be evident under all 

conditions. It is a little too anthropomorphic 

to expect young male pumas to avoid 

inbreeding under one situation and not 

another, especially since they don’t know if 

they are in Florida or Montana, i.e. they can 

only gain information about the number and 

distribution of potential mating opportunities 

by dispersing. Relative to the New Mexico 

data, based on our calculations of residency, 

we predict we should find cases of fathers 

not only mating with their daughters but 

could do so at least twice during their 

tenure. The data from Logan and Sweanor 

(2001) supported this prediction, thus 

refuting the inbreeding hypothesis. As 

fathers mating with daughters is genetically 

equivalent to sons mating with their 

mothers, we see no reason why a son would 

refuse to mate with his mother but then later 

mate with his daughter. It might be argued 

that a larger father could dominate over a 

smaller unreceptive daughter. However, we 

reject this argument because by the time 

young males reach dispersal age, they weigh 

equal to or more than their mothers 

(Laundré and Hernández 2002) and upon 

sexual maturity (21-27 months; Logan and

Sweanor 2001) could easily dominate over 

their smaller mother or sisters. As males do 

not seem reluctant to mate with close 

relatives, i.e. daughters, why don’t they wait 

around to mate with their sisters or mothers? 

We conclude that the only logical answer to 

this question is that the high level of conflict 

with their fathers or, by default, even more 

competitive older bigger new male arrivals 

(who have displaced their fathers), force 

young males to disperse out of their natal 

area at an age (13-15 months of age) before 

they are sexually mature (Sweanor 1990). 

Regarding prediction #3, the observed 

frequency distribution of HRDs was best fit 

by the competition model with a p(0) = 0 

and an increasing p(x) until a distance of 5 

HRDs. This model would reflect the effect 

of an increasing competitive ability of the 

young males as they disperse. In contrast, it 

is obvious that the data do not fit the most 

critical part of the inbreeding avoidance 

model: males should not settle within 2 

HRDs of their natal home range. 

Additionally, considering that even 5 HRDs 

may not be sufficient to reduce the chance of 

inbreeding with sisters (Shields 1987, 

Sweanor 1990), the high percent (>50%) of 

settlements within the first 4 HRDs argues 

strongly against inbreeding avoidance being 

the driving force behind male dispersal. 

The decreasing number of settlements 

between 5 and 7 HRDs likely reflects the 

fragmented nature of the 2 studies that 

contributed the majority of the data (54.2 %; 

Logan and Sweanor 2001 and this study). In 

both studies, the principal mountain range(s) 

of the study areas are separated from nearby 

larger mountain ranges (places young pumas 

are most likely to find vacant territories) by 

valleys 75 to 100 + km wide. Thus, there is 

a reduced likelihood of a male puma finding 

a territory in the 5-7 HRD range. 

Although neither model predicted the 

higher number of settlements we found at >9 

and up to 20 HRDs, this finding also does 



not support the inbreeding avoidance 

hypothesis. If a young male was dispersing 

primarily to avoid inbreeding, we would not 

predict such long distance dispersals where 

animals passed up what would appear to be 

ample suitable habitat in their dispersal 

movements. The only explanation for such 

long distance dispersals is they had to 

continue to travel because they were outcompeted 

by resident males or transients 

that had traveled further than themselves. 

Eventually, they would gain weight and 

experience enough to successfully compete 

for a territory (Sweanor 1990). 

Sweanor (1990) originally suggested that 

male residency times, the polygynous 

mating system, and male dispersal distances 

in pumas argued against inbreeding 

avoidance being the main driving force in 

dispersal of young male pumas. However, 

after presenting further evidence of 

aggression among male pumas and 

inbreeding between fathers and daughters, 

Logan and Sweanor (2001) concluded that 

inbreeding avoidance was likely the main 

driving force behind male puma dispersal. 

Their argument centered on the assumption 

that young males should only disperse far 

enough to “avoid” competition. We contend 

that with 15 - 18 % of puma populations 

being primarily transient males (Hemker et 

al. 1984, Ross and Jalkotzy 1992, 

Spreadbury et al. 1996, Laundré and Clark 

2003), no dispersing male ever has the 

luxury of avoiding competition for a 

territory. They concurred with the presence 

of these transients and observed that new 

males were constantly coming into their 

population and concluded from this that 

competition must be “tolerable”. They 

further argued that under the competition 

hypothesis, however, most of the male 

recruits in their growing population should 

have been offspring of the area. Because in 

their study most male recruits came from 

outside, they argued that this refuted the


competition hypothesis. It is unclear as to 

what they meant by “tolerable competition” 

but at any level of competition within its 

natal home range, a young male will be at a 

competitive disadvantage to his father or any 

new male immigrant; they will, by default, 

all be older and larger. Thus, under the 

competition hypothesis, we would expect no 

young males to compete successfully with 

their fathers or any older immigrant male 

that arrives to their home area. 

Consequently, we would predict that young 

males would all have to disperse, which is 

what all studies have found. Their chances 

of competing for a home range 1 HRD from 

their natal area is still low but greater than 

zero because they are now slightly older and 

bigger and we would predict that some, not 

many, might establish territories that close 

to their natal home range. This is supported 

by the data presented, 2 out of 48 dispersing 

males set up territories within the first HRD. 

The further they disperse in time and space, 

the more competitive they should be and we 

would predict that they would start winning 

the competition for territories. Thus rather 

than refuting the competition model, the fact 

that Logan and Sweanor (2001) found only 

6 males in 10 years born in the area setting 

up territories within the study area starting at 

≈ 2.5 HRDs, actually supports and would 

have been predicted by the competition 

hypothesis. Consequently, we contend that 

all the data presented by Logan and Sweanor 

(2001) concur with Sweanor’s (1990) 

original conclusion to reject the inbreeding 

hypothesis and provide strong support for 

the competition model. 

In conclusion, after our analysis, we also 

concur with Sweanor’s (1990) original 

assessment and rejected the inbreeding 

model because resident males do fight and 

kill their male offspring as well as younger 

transients, resident males do mate with their 

daughters, and dispersal distances were 

found to best fit that predicted by the 


competition model. We conclude that young 

males are forced out of their natal home 

range by their fathers or incoming males 

who, by default, will be older and stronger. 

We propose that they continue to disperse 

until they gain weight and experience 

enough to successfully takeover a territory. 

What we propose is that the avoidance of 

inbreeding is very likely not the driving 

force behind dispersal in young male pumas 

but rather, likely is a serendipitous 

consequence of dispersal driven by 

competitive interactions. 


The data from our study was gathered 

with the financial and logistic support of the 

following foundations and agencies: 

ALSAM Foundation, Boone and Crockett 

Club, Earthwatch Institute, Idaho State 

University, National Rifle Association, The 

Eppley Foundation, U.S. Bureau of Land 

Management, The Northern Rockies 

Conservation Cooperative, Idaho 

Department of Fish and Game, Mazamas, 

The Merril G. and Emita E. Hasting 

Foundation, Patagonia, Inc., SEACON of 

the Chicago Zoological Society, The 

William H. and Mattie Wattis Harris 

Foundation, and Utah Division of Wildlife. 

We would like to thank the many 

Earthwatch volunteers without whose help 

this work would not have been 

accomplished. We thank J. Linnell and M. 

Culver for their helpful comments on this 

manuscript. We also thank the various 

researchers whose published data we used in 

our analysis. It is through research efforts 

like theirs that we are able to advance our 

understanding of puma ecology and 

behavior. Lastly, we especially thank Kevin 

Allred and Ken Jafek. It is only through 

their tireless enthusiasm and willing use of 

their tracking dogs that our study was 

possible.


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LINDZEY, F.G., W.D. VAN SICKEL, B.B. 

ACHERMAN, D. BARNHURST, T.P. 

HEMKER, AND S.P. LAING. 1994. 

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mountain lion population in Wyoming. 


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conservation of an Enduring Carnivore. 

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panthers. National Geographic Research 

and Exploration 7:414-431. 

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ecology of the mountain lion (Felis 

concolor) of the Texas-Mexico border. 

Thesis, Sul Ross State University, 

Alpine, Texas, USA. 

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alternatives to the geometric distribution. 

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and inbreeding avoidance related? 

Animal Behaviour 32:94-112. 

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cougar (Puma concolor) in the northern 

Yellowstone ecosystem: Interactions 

with prey, bears, and humans. 

Dissertation. University of Idaho, 

Moscow, Idaho, USA.


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O.W. MORRIS. 1961. Notes on cougar 

productivity and life history. Journal of 

Mammalogy 42:204-217. 

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O’BRIEN. 1993. The consequences of 

demographic reduction and genetic 

depletion in the endangered Florida 

panther. Current Biology 3:340-350. 

ROSS, P.I. AND M.G. JALKOTZY. 1992. 

Characteristics of a hunted population of 

cougars in southwestern Alberta. 


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W.V. WILES, AND J.P. MESSICK. 1973. 

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systems: investigating their causal 

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Effects of Social Structure on Population 

Genetics. University of Chicago Press, 

Chicago, Illinois, USA. 

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with recommendations for management. 


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KAISNER, AND J. KOVAK. 1996. Cougar 

population characteristics in 

southeastern British Columbia. Journal 

of Wildlife Management. 60:962-969. 

SWEANOR, L.L. 1990. Mountain lion social 

organization in a desert environment. 

Thesis, University of Idaho, Moscow, 

Idaho, USA. 

SWEANOR, L.L., K.A. LOGAN, AND M.G. 

HORNOCKER. 2000. Cougar dispersal 

patterns, metapopulation dynamics and 

conservation. Conservation Biology 

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352.

COUGAR EXPLOITATION LEVELS AND LANDSCAPE CONFIGURATION: 

IMPLICATIONS FOR DEMOGRAPHIC STRUCTURE AND METAPOPULATION 

DYNAMICS 

DAVID C. STONER, Utah State University, Dept. of Forest, Range, and Wildlife Sciences, 

5230 Old Main Hill, Logan, UT, 84322-5230, USA, email: dstoner@cc.usu.edu 

MICHAEL L. WOLFE, Utah State University, Dept. of Forest, Range, and Wildlife Sciences, 

5230 Old Main Hill, Logan, UT, 84322-5230, USA, email: mlwolfe@cnr.usu.edu 

Abstract: Currently eleven states and two Canadian provinces utilize sport hunting as the 

primary mechanism for managing cougar (Puma concolor) populations. However the impacts of 

sustained harvest on demographic structure and population persistence are not well understood. 

Additionally, the range of non-biological factors influencing the rate of population recovery has 

not been thoroughly examined. We have been monitoring the cougar populations on Monroe 

Mountain in south-central Utah, and in the Oquirrh Mountains of north-central Utah since 1996 

and 1997, respectively. The critical management distinction between these sites is the degree of 

exploitation. The Monroe population is subjected to heavy annual hunting pressure and is 

characterized demographically by a younger age distribution, low survivorship, low fecundity, 

and declining density. In contrast, the population inhabiting the northeastern slope of the 

Oquirrhs is subjected to little or no hunting pressure and exhibits an older age distribution, 

relatively high survivorship and fecundity, a stable density, and a high emigration rate. Due in 

part to these differences, the Oquirrh and Monroe populations appear to exhibit source and sink 

dynamics within the regional metapopulation. Therefore the temporal scale of population 

recovery may depend on the interaction between the dominant harvest regime and the degree of 

landscape connectivity with neighboring patches. Aside from harvest, the interaction between 

patch configuration and anthropogenic fragmentation may be highly influential in the long-term 

prognosis for these populations. We discuss the implications of these demographic distinctions 

in light of enumeration uncertainties, habitat fragmentation, and landscape structure. 


161

162 

ASSESSING GPS RADIOTELEMETRY RELIABILITY IN COUGAR HABITAT 

TRISH GRISWOLD 

JAMES BRIGGS 

GARY KOEHLER 

STUDENTS AT CLE ELUM-ROSLYN SCHOOL DISTRICT, Cle Elum, Washington 

Abstract: Studies evaluating the effectiveness of GPS radiotelemetry have shown that the 

positional accuracy and rate of GPS fixes declines with increased forest canopy coverage 

(D’Eon, Serrouya, Smith, and Kochanny, 2002. Wildlife Society Bulletin, 30(2):430-439). Since 

GPS collars are being used to mark and monitor cougars (Puma concolor; Koehler and Nelson, 

7 th Mountain Lion Workshop), students, faculty, and volunteers at the Cle Elum-Roslyn Middle 

School, Washington, tested GPS location accuracy as part of Project CAT (Cougars and 

Teaching). We fitted domestic dogs (Canis familiaris) with the same GPS collars used to mark 

cougars and locational accuracy was measured in areas of known cougar habitat. GPS fixes were 

recorded and compared with UTM coordinates obtained from hand-held GPS receivers and 7.5minute 

topographic maps. Environmental factors, vegetation types, and physiographic 

parameters were recorded. It was felt that the dogs would closely approximate cougar movement 

patterns and give an index of reliability of GPS fixes for free-ranging cougars. While previous 

studies have addressed the reliability of GPS collar fixes, none have tested reliability of data 

collected in the rapidly suburbanizing ponderosa pine (Pinus ponderosa) and Douglas fir 

(Pseudosuga menziesii) forests of the eastern Cascade Mountains. This project gives the middle 

school students an opportunity to participate in the school-wide educational effort of cougar 

ecology. Students proposed and tested hypotheses and analyzed the data. 


USING GPS COLLARS TO DETERMINE COUGAR KILL RATES, ESTIMATE HOME 

RANGES, AND EXAMINE COUGAR-COUGAR INTERACTIONS 

POLLY C. BUOTTE, Research Assistant and GIS Specialist, Wildlife Conservation Society, 

2023 Stadium Dr. Suite 1A, Bozeman, MT 59030, USA, email: 

polly_thornton@hotmail.com 

TONI K. RUTH, Associate Conservation Scientist, Wildlife Conservation Society, 2023 Stadium 

Dr. Suite 1A, Bozeman, MT 59030, USA, email: truth@montanadsl.net 

Abstract: Single-species approaches to large carnivore conservation limits our understanding of 

carnivore assemblages and interactions at a community level and obtaining data on wide ranging, 

secretive species such as cougars and wolves can be particularly challenging. Beginning in 

2001, we collaborated with the Yellowstone Wolf Project and Interagency Grizzly Bear Team in 

applying GPS technology to examine patterns of resource use among cougars, wolves, and 

grizzly bears. In this paper, we address three topics relative to future analysis of GPS data on all 

three carnivores: 1) the efficacy of finding cougar-killed elk and deer carcasses through GPS 

locations, 2) differences in home range estimation from GPS versus VHF locations with 

implications to analysis of species overlap; and 3) interaction between two cougars with 

implications to addressing spatial-temporal interactions between cougars, wolves, and bears. We 

deployed store-on-board GPS collars (GPS Generation II, Telonics, Inc.) on two adult male 

cougars (M137 and M127) during the winter of 2001. Male M137’s collar acquired 612 GPS 

locations between Feb 11 and June 13, with a successful fix rate of 59.9%. Male M127’s collar 

acquired 370 GPS locations between Feb 27 and May 1, with a successful fix rate of 73.4%. 

Each collar was programmed to attempt a GPS fix every third hour, or eight times per day. We 

identified clusters of locations by calculating distance moved between consecutive GPS locations 

and by selecting groups of locations within 200 meters of each other. Identified clusters were 

located and searched in the field utilizing a hand-held GPS. For cougar M127, we additionally 

documented kill rate via intensive daily ground-based VHF telemetry sampling between March 5 

and April 10. Both ground and GPS methods yielded four kills during that time span. To 

examine differences in identification of home ranges we calculated home ranges using fixed 

kernel analysis. Male M137’s GPS data included a disjunct area of approximately 15 km 2 that 

was not identified from VHF locations. Preliminary analysis of interaction indicated two times 

when both cougars were at the same location, after which the subordinate male M127 moved 

away from the dominant male M137. During winter 2003, we deployed 5 Televilt Simplex GPS 

collars on cougars. The collars allow for remote downloads of data and are programmed to 

acquire locations simultaneous to locations of GPS collared wolves. Our goals during the next 

two years are to: 1) develop correction factors for both ground-based and GPS collected kill rates 

and, in collaboration with the Yellowstone Wolf Project and the Interagency Grizzly Bear Team 

to, 2) quantify spatial-temporal interactions between cougars, wolves and bears via subsequent 

moves analyses and utilize these data to develop a predictive model of carnivore movement and 

landscape use. 


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164 

FUNCTIONAL RESPONSE OF COUGARS AND PREY AVAILABILITY IN 

NORTHEASTERN WASHINGTON 

HILARY S. CRUICKSHANK, Large Carnivore Conservation Lab, Department of Natural 

Resource Sciences, Washington State University, PO Box 646410, Pullman, WA 99164, 

USA, email: hcruicks@mail.wsu.edu 

HUGH S. ROBINSON, Large Carnivore Conservation Lab, Department of Natural Resource 

Sciences, Washington State University, PO Box 646410, Pullman, WA 99164, USA, 

email: hsrobins@wsunix.wsu.edu 

CATHERINE LAMBERT, Large Carnivore Conservation Lab, Department of Natural Resource 


email: lambertcath@hotmail.com 

ROBERT B. WIELGUS, Large Carnivore Conservation Lab, Department of Natural Resource 


email: wielgus@mail.wsu.edu 

Abstract: Within the last ten years, a major change in the population structure of deer in western 

North America has taken place. Mule deer populations are sharply declining, while white-tailed 

deer populations are increasing. Researchers have suggested that cougar predation is a possible 

reason for the decline. The purpose of this project is to investigate cougar predation in a 

community where substantial populations of white-tailed deer, mule deer, and cougars overlap. 

We are testing two alternative hypotheses of cougar prey selection. H1, or the apparent selection 

hypothesis, states that equal selection by cougars for white-tailed deer and mule deer, but a 

higher reproductive rate by white-tailed deer is causing a decline in the mule deer population. 

H2 proposes that higher selection by cougars for mule deer is causing a decline in the population. 

Preliminary results suggest H2. The effect of predation on prey is determined by two factors: 1) 

functional response, and 2) prey availability. Functional response of cougars is quantified by the 

number of kills, per cougar, per unit time, and prey availability provides an estimate of the 

number and distribution of each prey species. The combination of these two factors may offer a 

more complete understanding of cougar prey selection. This research is in support of a larger 

study, which will use the apparent competition theory to examine alternative cougar management 

strategies. 


WHAT DOES TEN YEARS (1993-2002) OF MOUNTAIN LION OBSERVATION DATA 

REVEAL ABOUT MOUNTAIN LION–HUMAN INTERACTIONS WITHIN REDWOOD 

NATIONAL AND STATE PARKS? 

GREGORY W. HOLM, Wildlife Biologist, Redwood National Park, 219 Hilton Rd., Orick, CA 

95519, USA, email: Gregory_Holm@nps.gov 

Abstract: Mountain lions (Puma concolor) occur throughout Redwood National and State Parks 

(RNSP) and most other portions of northwest California. However, because they are not often 

observed, RNSP biologists have always been interested in recording mountain lion observations 

within the park. Prior to 4 mountain lion attacks in California between 1992-94, 2 resulting in 

fatalities, RNSP mountain lion observations were not compiled in a timely, consistent, or easily 

accessible manner. Since 1993, RNSP biologists have attempted to document and verify all 

mountain lion observations using a standard reporting form and database. Three hundred and 

seven mountain lion observations have been recorded within RNSP from 1993-2002 (mean ≅ 31; 

range 19-53). Most were observed between May and October during daylight hours, and 

involved a single mountain lion. While most observations (54%) involved a mountain lion near 

a road while people were in a vehicle, the remaining observations (46%) occurred while people 

were on trails or at other park facilities. The ultimate response of most mountain lions (68%) 

encountered on trails was to avoid humans, yet twenty percent of trail encounters involved some 

level of curiosity by the mountain lion towards humans. Although no human attacks were 

reported, there were 8 reports of aggressive behavior towards humans, 6 reports of following 

humans, and 1 report of a dog on a leash getting attacked. The observation data does not 

accurately reflect the actual distribution or timing (seasonally or daily) of mountain lion activity, 

and should be interpreted with caution due to inherent problems with observer experience, and 

report quality and verification. However, the information does allow managers to quickly 

identify when and where mountain lion-human interactions have occurred, and more effectively 

focus management actions to prevent or reduce future mountain lion-human interactions within 

RNSP. 


165

166 

DEPREDATION TRENDS IN CALIFORNIA 

SARAH REED, Mountain Lion Foundation, PO Box 1896, Sacramento, CA 95814, USA 





Abstract: Since 1972 more than 1,600 California mountain lions have been killed under 

depredation permits. The number of lions killed annually has increased, with a peak of 149 lions 

killed in 2000. Although some permits are issued for losses incurred by traditional, 

economically viable, open range livestock operations, incidents on ranchettes and "hobby farms" 

are increasing. We evaluate trends in depredation permitting, including analysis of depredation 

events geographically, by parcel size and size of herd, and relative to human population and 

development trends. We conclude with new approaches to mountain lion conservation that stress 

science to establish a factual basis for dialogue, community involvement, and developing 

partnerships with diverse stakeholders. 


THE DISTRIBUTION OF PERCEIVED ENCOUNTERS WITH NON-NATIVE CATS IN 

SOUTH AND WEST WALES, UK: RELATIONSHIP TO MODELED HABITAT 

SUITABILITY 

A.B. SMITH, Exotic Cat Group, Geography Department, University of Wales Swansea, 

Singleton Park, Swansea SA2 8PP, UK, email: exoticcatproject@swan.ac.uk 

F.A. STREET PERROTT, Exotic Cat Group, Geography Department, University of Wales 

Swansea, Singleton Park, Swansea SA2 8PP, UK 

T. HOOPER, Exotic Animals Register, 85 Risedale Road, Ashton Vale, Bristol BS3 2RB, UK 

Abstract: Reports of perceived encounters with exotic cats in the British countryside have 

greatly increased in recent years. The species described (notably melanistic leopards, pumas and 

lynxes) were widely bred in the UK prior to the 1976 Dangerous Wild Animals Act, and do not 

correspond to those most familiar to the general public (such as lions, tigers and cheetahs). 

Some of them are still being illegally imported or reared for ‘canned hunts’. Following the 

recent discovery of leopard tracks in West Wales, and calls for action by Members of the UK 

Parliament and the Welsh National Assembly, the Welsh Agriculture Department has officially 

begun to collect statistics on sightings and livestock kills. In this independent study, we have 

analysed a database of 170 georeferenced encounter reports obtained from the police, news 

media and members of the public. In the absence of confirmatory DNA, physical or photographic 

evidence, encounter reports require very careful screening for reliability, based on the 

characteristics of the witness(es); the validity of the identification, taking into account perceived 

cat size and shape (morphotype) and behaviour; indicators of scale, distance and lighting 

conditions; and the suitability of the habitat. The distributions of potential habitats within South 

and West Wales have been modelled with a GIS using standard habitat characteristics, such as 

prey-species presence, disturbance levels, geomorphology and land-use data. The spatial pattern 

of encounters does not show the clustering that might be expected if they represent a purely 

sociological phenomenon. Instead, the distributions of specific morphotypes appear to be closely 

related to the degree of habitat suitability, thereby strengthening the case for the presence of nonnative 

cat species in the UK. 


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168 

PUMA ACTIVITY AND MOVEMENTS IN A HUMAN-DOMINATED LANDSCAPE: 

CUYAMACA RANCHO STATE PARK AND ADJACENT LANDS IN SOUTHERN 

CALIFORNIA 

LINDA L. SWEANOR, Wildlife Health Center, University of California - Davis, Southern 

California Puma Project Field Station: P.O. Box 1114, Julian, CA 92036, USA, email: 


KENNETH A. LOGAN, Wildlife Health Center, University of California - Davis, Southern 

California Puma Project Field Station: P.O. Box 1114, Julian, CA 92036, USA, email: 

klogan2@mindspring.com 

JIM W. BAUER, Wildlife Health Center, University of California - Davis, Southern California 

Puma Project Field Station, P. O. Box 1203, Julian, CA 92036, USA, email: 


WALTER M. BOYCE, Wildlife Health Center, University of California - Davis, One Shields 

Avenue, Davis, CA 95616, USA, email: wmboyce@ucdavis.edu 

Abstract: Although puma attacks are exceedingly rare, statistics indicate dangerous encounters 

between humans and pumas are on the rise. In California there have been 7 verified puma attacks 

resulting in 2 human deaths during the last 10 years; 2 of those attacks and 1 death occurred in 

Cuyamaca Rancho State Park (CRSP). Because of the high number of reported puma sightings 

in CRSP each year (range = 18-50 from 1993-2002) and the large, increasing number of human 

visitors (over 500,000 people visited the 50-square-mile park in 2001), park authorities were 

concerned about the potential for further dangerous puma-human encounters. A study was 

initiated in January 2001 to understand puma behavior relative to human activity, to help 

minimize conflicts between pumas and humans, and to assist the development of long-term 

conservation strategies for pumas in the CRSP area. Specific objectives of the project were to: 

determine the number and characteristics of pumas using CRSP; map puma home ranges and 

determine important puma habitats and their juxtaposition relative to human use areas; examine 

puma movements (e.g., daily, seasonal) relative to areas of human activity (e.g., trails, roads, 

campgrounds); examine puma diet to determine what prey species are most important as puma 

food and to what extent, if any, domestic animals contribute to their diet; and use the data to 

formulate management recommendations. To obtain information on puma home ranges, 

movements and behavior, and to find prey killed by pumas, independent and adult pumas are 

being captured and fitted with Televilt GPS collars. As of March 2003, 11 pumas (6 adult males, 

5 adult females) had been captured in and around CRSP and fitted with GPS collars. To date, 

collars have yielded over 6400 locations. Additionally, human use of trails is being measured 

seasonally by placing infrared counters (TrailMaster monitors) along 4 trail systems within 

CRSP. This paper reports on project progress to date. 


MODELING OFFSPRING SEX RATIOS AND GROWTH OF COUGARS 

DIANA M. GHIKAS, Department of Biological Sciences, University of Calgary, 2500 

University Drive N.W., Calgary, AB T2N 1N4, Canada, email: dghikas@ucalgary.ca 

MARTIN JALKOTZY, Arc Wildlife Services Ltd., 3527-35th Avenue S.W., Calgary, AB T3E 

1A2, Canada, email: martin_jalkotzy@nucleus.com 

IAN ROSS, Arc Wildlife Services Ltd., 3527-35th Avenue S.W., Calgary, AB T3E 1A2, Canada 

RALPH SCHMIDT, Arc Wildlife Services Ltd., 3527-35th Avenue S.W., Calgary, AB T3E 

1A2, Canada 

SHANE A. RICHARDS, Department of Biological Sciences, University of Calgary, 2500 

University Drive N.W., Calgary, AB T2N 1N4, Canada 

Abstract: We further examined data from the Sheep River cougar study conducted in southwest 

Alberta from 1981-94. We asked is there evidence of an equal or constant offspring sex ratio, or 

do sex ratios vary over time, as a function of the mother's age, geographic location or population 

size. Logan and Sweanor (2001) analysed offspring sex ratios as a function of mother's age and 

found that sex ratios of first litters were significantly different from subsequent litters and 1:1. 

Logan and Sweanor (2001) suggested that offspring sex ratios might be influenced by the 

mother's physical state (i.e., young mothers produce less-costly females so energy can be 

allocated to growth). We investigated possible relationships between offspring sex ratios and 

cougar growth, and whether growth varied by sex and geographic location. The study area was 

divided into east and west locations based coarsely on prey abundance and cougar mortality. 

Probabilistic models were formulated for the sex-ratio analysis. A deterministic model based on 

the flexible Richards curve (Maehr and Moore 1992) was used to predict mass growth. Model 

predictions that were the most parsimonious with the data were identified using corrected Akaike 

Information Criterion. Parameters were estimated using maximum log-likelihood. The most 

parsimonious model predicted that offspring sex ratios vary yearly. Evidence was not strong for 

sex ratios varying as a linear function of mother's age. The growth model predicted that females 

attain 91-92 % of adult mass by 25-26 mos, indicating that growth is largely completed prior to 

first reproduction ( x =30.0 ± 1.8 mos SE, Ross and Jalkotzy 1992). In years with poor resource 

conditions, the mother's physical state may result in more female offspring being reared than 

male. Predicted mean mass at age of independence for male and female offspring was 48.8 kg 

and 34.1 kg, respectively, inferring males are more costly to rear. The growth model that varied 

by sex only had the highest weight of evidence; adding geographic location did not result in a 

more parsimonious model. All growth models were unable to accurately estimate birth mass, 

which was also found by Maehr and Moore (1992). 


Logan, K. A., and L.L. Sweanor. 2001. Desert Puma: Evolutionary ecology and conservation of an enduring carnivore. Island 

Press, Washington. 

Maehr, D.S., and C.T. Moore. 1992. Models of mass growth for 3 North American cougar populations. J. Wildl. Manage. 56: 

700-707. 

Ross, P.I., and M.G. Jalkotzy. 1992. Characteristics of a hunted population of cougars in southwestern Alberta. J. Wildl. 

Manage. 56: 417-426. 


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170 

MOUNTAIN LION SURVEY TECHNIQUES IN NORTHERN IDAHO: A THREE-FOLD 

APPROACH 

CRAIG G. WHITE, Idaho Department of Fish and Game, 1540 Warner Avenue, Lewiston, ID 

83501, USA, email: cwhite@idfg.state.id.us 

PETER ZAGER, Idaho Department of Fish and Game, 1540 Warner Avenue, Lewiston, ID 

83501, USA, email: pzager@idfg.stat.id.us 

LISETTE WAITS, Department of Fish and Wildlife Resources, University of Idaho, Moscow, 

ID 83843, USA, email: lwaits@uidaho.edu 

Abstract: Management of mountain lions (Puma concolor) in Idaho relies largely on harvest 

data. This type of data is limited in scope and relays little information to the manager regarding 

population trend or density. Intensive radio telemetry studies involving capture and recapture 

can provide an estimation of density but are expensive. Currently researchers are exploring 

techniques to index or estimate population size by identifying individuals by their DNA. We 

outline three different techniques to “capture” and “recapture” mountain lion hair and/or tissue 

for DNA analysis: biopsy darts, rub tree stations, and legally harvested lions. Techniques are 

being implemented on two study areas in north-central Idaho, the Lochsa/North Fork of the 

Clearwater River and the South Fork of the Clearwater River. Efforts by both volunteer 

houndsmen and hired houndsmen over 1½ lion harvest seasons have resulted in ≥15 DNA 

samples from lion treed and released. Over the same period ≥15 DNA samples have been turned 

in from legally harvested lions. In 2002, we placed 51 rub stations and recorded 42 visits over 

the 3 sampling periods. Seventeen of the visits resulted in ≥1 hair. Preliminary results indicate 

that 1 visit was from a lion, 7 visits were by bear, and 7 visits were possibly a lion or bear, and 2 

visits by other species. Improvements to the techniques are ongoing. This study will allow us to 

identify individuals in the mountain lion populations within our study areas and thus obtain a 

minimum population size. Number of captures each year will serve as an index of population 

trend. We will also explore the use of capture-recapture modeling to estimate population sizes. 

Our approach attempts to limit resources expended in capturing and marking animals, while still 

providing an index and potential population estimate within our study areas. 


MOUNTAIN LIONS IN SOUTH DAKOTA: RESULTS OF A 2002 PUBLIC OPINION 

SURVEY 

LARRY M. GIGLIOTTI, Planning Coordinator/Human Dimensions Specialist, South Dakota 

Department of Game, Fish and Parks, 523 E. Capitol, Pierre, SD 57501, USA, email: 

larry.gigliotti@state.sd.us 

DOROTHY M. FECSKE, Department of Wildlife and Fisheries Sciences, Box 2140B, South 

Dakota State University, Brookings, SD 57007, USA, email: gdf@rapidnet.com 

JONATHAN A. JENKS, Department of Wildlife and Fisheries Sciences, Box 2140B, South 

Dakota State University, Brookings, SD 57007, USA, email: 

Jonathan_Jenks@sdstate.edu 

Abstract: Mountain lions (Puma concolor) are a state threatened species in South Dakota, 

although there is an established breeding population in the Black Hills. The Department of 

Game, Fish and Parks (GFP) is currently funding a multi-year research project through South 

Dakota State University to learn more about the status of mountain lions. The information will 

be used by GFP to develop a mountain lion management plan. Public opinion and understanding 

of mountain lions will be a critical component for developing and implementing a management 

plan. This public opinion survey was the first step in developing the social component (human 

dimensions) of the plan. The survey was conducted in the early spring of 2002. Of 1,783 

deliverable questionnaires mailed to South Dakota residents, 1,114 usable questionnaires were 

returned for a total return rate of 62.57%. A one-page survey of non-respondents also was 

conducted; of those, 103 (19.5%) were returned. Overall, the majority of respondents (>50%) 

believed that presence of mountain lions was an indication of a healthy environment, lions and 

hunters did not compete for deer, if people modified a few behaviors they could coexist with 

lions, and lions should be able to exist wherever they occurred in South Dakota. Survey results 

were used to develop an attitude model to provide a framework for understanding public opinion 

of mountain lions. The model was intuitive, but derived empirically using a cluster analysis 

procedure from respondents’ answers to 12 questions. The model represented a continuum of 

attitudes ranging from strongly supportive of to strongly disliking mountain lions. Based on the 

model, 22.7% of the respondents were strongly pro-lion, 33.7% slightly pro-lion, 11.3% neutral, 

22.5% slightly contra-lion, and 9.8% of the respondents strongly contra-lion. Cluster names 

were descriptive of the general attitudes held toward mountain lions in South Dakota, and 

responses provided to other questions in the survey were used to further describe each clustergroup. 


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CRITICAL COUGAR CROSSING AND BAY AREA REGIONAL PLANNING 

MICHELE KORPOS, Live Oak Associates, Inc., 6830 Via Del Oro, Suite 205, San Jose, CA 

95119, USA, email: mkorpos@loainc.com 

Abstract: The goal of this presentation is to illustrate the need for habitat conservation and 

regional planning on behalf of cougars (Puma concolor), which play an integral role in the health 

of their ecosystems. As a keystone species, “further degradation of [cougar] habitat connectivity 

will lead to cascading impacts down through successively lower trophic levels…” (Jigour 2000). 

County borders are human constructs with no ecological relevance. It can be assumed certain 

cougar home ranges overlap San Mateo, Santa Clara and Santa Cruz Counties (California), while 

others may overlap Monterey, San Benito and Santa Cruz Counties. These counties contain the 

Santa Cruz Mountains, and the Gabilan and Diablo Mountain Ranges. The challenge is to 

maintain land connections between large patches of intact habitat through open communication 

among county agencies and through regional planning efforts. 

By developing land on a project-by-project basis, counties promote habitat fragmentation. Left 

unchecked, human development in and around the Santa Cruz Mountains will continue to 

fragment cougar habitat, leading to geographic isolation, and the eventual demise of our local 

population. Maintaining large tracts of land and providing connections through less hospitable 

landscapes are critical to ensuring the future health of cougar populations and the wildlife that 

share their ecosystems. 


Jigour, V. 2000. Correspondence to Rusty Areias, Director, Department of Parks and Recreation, dated 31 January 2000. 


Holly Akenson 

Taylor Ranch Field Station, Univ. of Idaho 

HC 83 Box 8070 

Cascade, ID 83611 

USA 

888-842-7547 

tayranch@direcpc.com 

Jim Akenson 

Taylor Ranch Field Station, Univ. of Idaho 

HC 83 Box 8070 

Cascade, ID 83611 

USA 

888-842-7547 

tayranch@direcpc.com 

Kevin Allred 

373 East 300 South 

Burley, ID 83318 

USA 

208-678-3681 

Chuck Anderson 

Wyoming Coop. Fish and Wildlife Research Unit 

Box 3166, University Station 

Laramie, WY 82071 

USA 

307-766-2091 

cander@uwyo.edu 

Jerry Apker 

Colorado Division of Wildlife 

0722 South Road 1 East 

Monte Vista, CO 81144 

USA 

719-587-6922 

jerry.apker@state.co.us 

Fred Armstrong 

Guadalupe Mountain National Park 

HC 60 Box 400 

Salt Flat, TX 79847 

USA 

915-828-3251 

Fred_Armstrong@nps.gov 

LIST OF PARTICIPANTS 


LIST OF PARTICIPANTS 173 

Matt Austin 

BC Ministry of Water, Land, and Air Protection 

PO Box 9374 

Victoria, BC V8V 1N4 

Canada 

250-387-9799 

Matt.Austin@gems7.gov.bc.ca 

Dave Avey 


3030 Energy Lane, Suite 100 

Casper, WY 82604 

USA 

307-473-3420 

Dave.Avey@wgf.state.wy.us 

David Baron 

2338 18 th Street 

Boulder, CO 80304 

USA 

303-443-2341 

dbaron@aya.yale.edu 

Jim Bauer 

Univ. of California-Davis, Wildlife Health Center 

PO Box 1203 

Julian, CA 92036 

USA 

760-767-4331 


Rich Beausoleil 


3860 Chelan Hwy 

Wenatchee, WA 98801 

USA 

509-679-3858 

beausrab@dfw.wa.gov 

Scott Becker 



Lander, WY 82520 

USA 

307-332-2688 

scott.becker@wgf.state.wy.us

174 LIST OF PARTICIPANTS 

Chris Beldon 

Florida Fish and Wildlife Conservation Commission 

4005 South Main Street 

Gainesville, FL 32601 

USA 

352-955-2230 

Chris.Belden@fwc.state.fl.us 

William Betty 

Eastern Puma Research Network 

49B Punch Bowl Trail 

West Kingston, RI 02892 

USA 

401-789-4026 

wjbetty@peoplepc.com 

Roman Biek 

Fish and Wildlife Biology Program 

University of Montana 

Missoula, MT 59812 

USA 

406-243-6193 

rbiek@selway.umt.edu 

Mario Biondini 

Department of Animal and Range Sciences 

North Dakota State University 

Fargo, ND 58105 

USA 

701-231-8208 

mario.biondini@ndsu.nodak.edu 

Dan Bjornlie 




USA 

307-332-2688 

dan.bjornlie@wgf.state.wy.us 

H. Webb Blessley 

The Cougar Fund 

910 Baja 

Laguna Beach, CA 92651 

USA 

702-275-7820 

wblessley@aol.com 

Steven Bobzien 

East Bay Regional Park District 

4422 Rockwood Avenue 

Napa, CA 94558 

USA 

510-544-2347 

sbobzien@ebparks.org 

Roger Bredehoft 


1017 Reynolds 


USA 

307-745-4402 

rbredeho@wyoming.com 

Jim Briggs 

Cle Elum-Roslyn School District - Project CAT 

2694 SR 903 

Cle Elum, WA 98922 

USA 

Doug Brimeyer 


PO Box 67 

Jackson, WY 83001 

USA 

307-733-2321 

Doug.Brimeyer@wgf.state.wy.us 

Lara Brongo 

Auburn University 

1925 Trexler Court 

Raleigh, NC 27606 

USA 

334-332-1124 

llbrongo@yahoo.com 

Howard Buffet 

Howard G. Buffet Foundation 

PO Box 4508 

Decatur, IL 62521 

USA 

217-429-3988 

hgbfoundation@yahoo.com 

Polly Buotte 

Wildlife Conservation Society 

Box 299 

Gardiner, MT 59030 

USA 

406-848-7683 

polly_thornton@hotmail.com 

Clint Cabanero 

South Coast Wildlands Project 

PO Box 2493 

Monrovia, CA 91016 

USA 

626-599-9585 

clint@scwildlands.org 


Steve Cain 

Grand Teton National Park 

PO Box 170 

Moose, WY 83012 

USA 

307-739-3485 

Steve_Cain@nps.gov 

Franz Camenzid 

Jackson Hole Conservation Alliance 

PO Box 2835 


USA 

307-733-9417 

Franz@jhalliance.org 

David Choate 

University of Notre Dame 

107 Galvin Life Science Center 

Notre Dame, IN 46556 

USA 

574-631-0949 

dchoate@nd.edu 

Stacy Courville 

CSKT – Wildlife Management Program 

PO Box 278 

Pablo, MT 59855 

USA 

406-883-2888 

stacyac@cskt.org 

Hilary Cruickshank 

Washington State University 

PO Box 651 

Rossland, BC V0G 1Y0 

Canada 

250-362-3310 

hcruicks@mail.wsu.edu 

Michelle Cullens 

Mountain Lion Foundation 

PO Box 1896 

Sacramento, CA 95812 

USA 

916-442-2666 

cullens@mountainlion.org 

Chris Daubin 


1205 Mary Anne 

Riverton, WY 82501 

USA 

307-856-4982 

cdaubin@tcinc.net 

Jeff Davis 

Wildlife Services 

PO Box 131 

Olancha, CA 93549 

USA 

760-937-6788 

Deanna Dawn 

University of California – Davis 

20231 Blauer Drive 

Saratoga, CA 95070 

USA 

408-741-5156 

Rich DeSimone 

Montana Fish, Wildlife, and Parks 

1420 East 6 th Avenue 

Helena, MT 59620 

USA 

406-444-0358 

rdesimone@state.mt.us 



Leonard Two Eagle 

Rosebud Sioux Tribe Game, Fish, and Parks 

PO Box 291 

Parmelee, SD 57566 

USA 

605-747-2289 

ln2eagle@hotmail.com 

Dorothy Fecske 

South Dakota State University 

Box 2140B 

Brookings, SD 57007 

USA 

605-688-6121 

gdf@rapidnet.com 

Lee Fitzhugh 


One Shields Drive 

Davis, CA 95616 

USA 

530-752-1496 

elfitzhugh@ucdavis.com 

Flora Fitzhugh 

913 Purdue Drive 

Woodland, CA 95695 

USA 

530-668-1138


Steve Fitzwater 

National Trappers Association 

PO Box 106 

Dubois, ID 83423 

USA 

208-745-6664 

dubois5@yahoo.com 

Gary Fralick 


PO Box 1022 

Thayne, WY 83127 

USA 

307-883-2998 

gfralick@silverstar.com 

Steve Galentine 

U.S. Department of Agriculture 

29469 East Vine Avenue 

Escalon, CA 95320 

USA 

209-605-4934 

sgalentine@mindspring.com 

Emily Garding 

Grand Canyon National Park 

Box 129 

Grand Canyon, AZ 86023 

USA 

928-638-7648 

Emily_Garding@nps.gov 

Jacquie Gerads 

North Dakota Game and Fish Department 

100 North Bismarck Expressway 

Bismarck, ND 58501 

USA 

701-328-6613 

jgerads@state.nd.us 

Diana Ghikas 

University of Calgary 

2500 University Drive NW 

Calgary, AB T3E 1A2 

Canada 

403-299-2797 

dghikas@ucalgary.ca 

Glenn Gibbons 

Oglala Sioux Parks and Recreation Authority 

PO Box 570 

Kyle, SD 57752 

USA 

605-455-2584 

aranger@gwtc.net 

Larry Gilbertson 

Nevada Division of Wildlife 

#60 Youth Center Road 

Elko, NV 89801 

USA 

775-777-2302 


Carlos Lopez Gonzalez 

Escuela de Biologia-Facultad de Ciencias Naturales 

Universidad Autonoma de Queretaro 

Apdo. Postal 184 

Queretaro, Queretaro 76010 

Mexico 

442-215-4777 

Cats4mex@aol.com 

Chelsea Gordon 

Cle Elum-Roslyn School District – Project CAT 

2694 SR 903 


USA 

Kevin Grady 

Boulder County Open Space 

1933 Geer Canyon Drive 


USA 

720-406-9178 

Kgrady@co.boulder.co.us 

Steve Griffin 

South Dakota Department of Game, Fish, and Parks 

3305 West South Street 

Rapid City, SD 57702 

USA 

605-394-2391 

steve.griffin@state.sd.us 

Torey Griswold 


2694 SR 903 


USA 

Trish Griswold 


2694 SR 903 


USA

Carolyn Grygiel 

Natural Resources Management 

North Dakota State University 

Fargo, ND 58105 

USA 

701-231-8180 

carolyn.grygiel@ndsu.nodak.edu 

Kerry Gyekis 

Writer/Private Consultant Forester 

RR 1, Box 213 

Morris, PA 16938 

USA 

570-723-8251 

gyekis@epix.net 

Keto Gyekis 

107 Woods Trail 

Delton, MI 49046 

269-623-3240 

keto@uplink.net 

Jon Hanna 

Arizona Game and Fish Department 

7200 E. University 

Mesa, AZ 85207 

USA 

480-981-9400 

Kevin Hansen 

Yosemite National Park 

PO Box 434 

Yosemite, CA 95389 

USA 

209-372-8870 

Jan V. Hart 

USGS – Southwest Biological Research Center 

PO Box 5614, NAU 

Flagstaff, AZ 86011 

USA 

928-556-7466 

Jan.Hart@nau.edu 

John Hart 

Hartwood Natural Resource Consultants 

1390 Curt Gowdy Drive 

Cheyenne, WY 82009 

USA 

307-778-3993 

huntwyoming@aol.com 

Dee Dee Hawk 

Wyoming Game and Fish Lab 

PO Box 3312, University Station 


USA 

307-766-6313 

Dhawk@uwyo.edu 

Lisa Haynes 

University of Arizona 

133 West 2 nd Street, Rear 

Tucson, AZ 85705 

USA 

520-320-1841 

lynxrufus@earthlink.net 

Lucina Hernández 

Instituto de Ecologia, A.C. 

Km 5 a Carr. de Mazatlán s/n 

Durango, Dgo C.P 34100 

México 

+52-618-812-1483 

lucina@fauna.edu.mx 

Greg Hiatt 


PO Box 186 

Sinclair, WY 82334 

USA 

307-324-7927 

gshiatt@trib.com 



Ryan Hill 


2694 SR 903 


USA 

Mary Hindelang 

Michigan Tech University 

40032 N. Lower Worham Road 

Chassell, MI 49916 

USA 

906-523-4014 

mlhindel@mtu.edu 

Dave Hoerath 

Boulder County Parks and Open Space 

5305 Spine Road, Unit B 


USA 

303-516-9364 

dhoerath@co.boulder.co.us


Mollie Hogan 

The Nature of Wildlworks 

PO Box 109 

Topanga, CA 90290 

USA 

310-455-0550 

wildworks1@aol.com 

Greg Holm 

Redwood National Park 

PO Box 7 

Orick, CA 95555 

USA 

707-464-6101 

gregory_holm@nps.gov 

Bernie Holz 


PO Box 850 

Pinedale, WY 82941 

USA 

307-367-4353 

Bernie.Holz@wgf.state.wy.us 

Mike Hooker 




USA 

307-332-2688 

michael.hooker@wgf.state.wy.us 

Rick Hopkins 

Live Oak Associates, Inc. 

6830 Via del Oro, Suite 205 

San Jose, CA 95119 

USA 

408-281-5885 

rhopkins@loainc.net 

Betsy Howell 

525 Benton 

Port Townsend, WA 98368 

USA 

360-379-0582 

betsyhowell@olympus.net 

Neil Hymas 


Box 368 

Cokeville, WY 83114 

USA 

307-279-3466 

nhymas@allwest.net 

Rose Jaffe 

Montana Fish, Wildlife, and Parks 

PO Box 200701 

Helena, MT 59620-0701 

USA 

406-444-1276 

rjaffe@state.mt.us 

Lynn Jahnke 


PO Box 6249 

Sheridan, WY 82801 

USA 

307-672-7418 

Lynn.Jahnke@wgf.state.wy.us 

Martin Jalkotzy 

Arc Wildlife Services Ltd. 

3527 – 35 Ave SW 


Canada 

403-240-3361 

martin_jalkotzy@nucleus.com 

Jesse Millen-Johnson 

Bates College 

Box 486 

Lewiston, ME 04240 

USA 

207-674-2927 

jmillenj@bates.edu 

Don Jones 

Zion National Park 

State Route 9 

Springdale, UT 84767 

USA 

435-772-0212 

don_jones@nps.gov 

Tom Keegan 

Idaho Department of Fish and Game 

PO Box 1336 

Salmon, ID 83467 

USA 

208-756-2271 

tkeegan@idfg.state.id.us 

Sean Kelly 

Utah Division of Wildlife Resources 

PO Box 606 

Cedar City, UT 84720 

USA 

435-691-5701 

seankelly@utah.gov 


Marc Kenyon 


7548 Event Way 


USA 

916-320-6654 

mwkenyon@ucdavis.edu 

Heather Keough 

Utah State University 

6991 S. Hwy 165 

Hyrum, UT 84319 

USA 

435-881-2856 

heatherkeough@hotmail.com 

Brian Kertson 

Washington Coop. Fish and Wildlife Research Unit 

23427 NE 28 th PL 

Sammamish, WA 98074 

USA 

425-941-0278 

gulogulo@hotmail.com 

Sharon Kim 

Zion National Park 

State Route 9 

Springdale, UT 84767 

USA 

435-772-0212 

sharon_kim@nps.gov 

Mike Kintigh 

South Dakota Game, Fish, and Parks 

3305 W. South Street 

Rapid City, SD 57702 

USA 

605-394-2391 

Mike.Kintigh@state.sd.us 

Gary Koehler 


PO Box 102 


USA 

509-260-0477 

gmkmmk@cleelum.com 

Andrea Kortello 

University of Idaho 

PO Box 4297 

Banff, AB T1L 1E7 

Canada 

403-762-5339 

kortello@yahoo.com 

Caroline Krumm 

CSU – Colorado Division of Wildlife 

317 W. Prospect 

Fort Collins, CO 80524 

USA 

970-215-3759 

ckrumm@yahoo.com 



Pamela Kyselka 

Navajo Nation – Department of Fish and Wildlife 

PO Box 1480 

Window Rock, AZ 86515 

USA 

928-871-6451 

p_kyselka@hotmail.com 

Carl Lackey 


1060 Mallory Wy. 

Carson City, NV 89701 

USA 

775-720-6130 

cdembears@aol.com 

Catherine Lambert 

Washington State University 

1712 NW Lamont St. 

Pullman, WA 99163 

USA 

509-335-4084 

lambertcath@wsu.edu 

Melanie Lambert 

The Summerlee Foundation 

716 W. Tejon, Suite 9 

Colorado Springs, CO 80903 

USA 

800-256-7515 

mal3@summerlee.org 

Darrell Land 


566 Commercial Blvd. 

Naples, FL 34104-4709 

USA 

239-643-4220 

Darrell.Land@fwc.state.fl.us 

John Laundré 

Instituto de Ecologia, A.C. 

Km 5 a Carr. de Mazatlán s/n 

Durango, Dgo C.P 34100 

México 

+52-618-812-1483 

launjohn@prodigy.net.mx


Cheryl Le Drew 

Lotek Wireless Inc. 

115 Pony Drive 

Newmarket, ON L3Y 7B5 

Canada 

905-836-6680 

cledrew@lotek.com 

Fred Lindzey 

Wyoming Coop. Fish and Wildlife Research Unit 

Box 3166, University Station 


USA 

307-766-5415 

flindzey@uwyo.edu 

Kenneth Logan 


2300 South Townsend Avenue 

Montrose, CO 81401 

USA 

970-252-6013 

Ken.Logan@state.co.us 

Mark Lotz 


566 Commercial Blvd. 

Naples, FL 34104-4709 

USA 

239-735-0773 

Mark.Lotz@fwc.state.fl.us 

Cara Blessley Lowe 


Box 122 


USA 

310-562-4021 

cara@mangelson.com 

Marissa Luchau 


2694 SR 903 


USA 

Lisa Lyren 

US Geological Survey 

1147 E. 6 th Street 

Corona, CA 92879 

USA 

909-735-0773 

llyren@usgs.gov 

Tom Mangelson 


Box 122 


USA 

307-733-6179 

tom@mangelson.com 

Marcie Maras 


2694 SR 903 


USA 

Beth Marker 


2694 SR 903 


USA 

markerb@cleelum.wednet.edu 

Gary Matson 

Matson’s Lab 

PO Box 308 

Milltown, MT 59851 

USA 

406-258-6286 

gjmatson@montana.com 

David Mattson 

USGS – Southwest Biological Research Center 

PO Box 5614, NAU 

Flagstaff, AZ 86011 

USA 

928-556-7466 

David.Mattson@nau.edu 

Roy McBride 

Livestock Protection Co. 

Box 178 

Ochopee, FL 34141 

USA 

239-695-2287 

Helen McGinnis 

Eastern Cougar Foundation 

PO Box 300 

Harman, WV 26270 

USA 

304-227-4166 

helenmcginnis@meer.net 


Craig McLaughlin 


1594 W. North Temple 

Salt Lake City, UT 84114 

USA 

801-538-4758 

craigmclaughlin@utah.gov 

Daryl Meints 


1515 Lincoln Road 

Idaho Falls, ID 83401 

USA 

208-525-7290 

dmeints@idfg.state.id.us 

Stephanie Middlebrooks 

Rosebud Sioux Tribe 

Box 300 

Rosebud, SD 57570 

USA 

605-747-2289 

middlebrooks@post.com 

Michael Middleton 

Muckleshoot Indian Tribe 

39015 172 nd Ave SE 

Auburn, WA 98092 

USA 

360-802-2202 

mikem@mitwildlifeculture.org 

Jessica Montag 

University of Montana 

2224 West Sussex 

Missoula, MT 59812 

USA 

406-243-6611 

jmontag@selway.umt.edu 

Dave Moody 




USA 

307-332-2688 

dave.moody@wgf.state.wy.us 

Shane Moore 

Box 2980 


USA 

307-733-8862 

moorefilms@wyoming.com 

Don Morgan 

Southern Hills Animal Clinic 

Box 67 

Pringle, SD 57773 

USA 

605-673-3503 

Susan Morse 

Keeping Track Inc. 

Wolfrun 55A Bentley Lane 

Jericho, VT 05465 

USA 

Kerry Murphy 

Yellowstone National Park 

PO Box 168 

Yellowstone National Park, WY 82190 

USA 

307-344-2393 

kerry_murphy@nps.gov 

Steve Nadeau 


600 South Walnut St., Box 25 

Boise, ID 83707 

USA 

208-334-2920 

snadeau@idfg.state.id.us 

Sharon Negri 

Wild Futures 

353 Wallace Way 

Bainbridge Island, WA 98110 

USA 

206-780-9718 

snegri@igc.org 



Evelyn Nelson 


2694 SR 903 


USA 

Mark Nelson 


500 Western Hills Blvd. 


USA 

307-638-8354 

mnelson@wyoming.com


Ryan Nelson 


2694 SR 903 


USA 

Jesse Newby 


Box 299 


USA 

406-848-7683 

newby_jr@yahoo.com 

M. Cathy Nowack 

Cat Tracks Wildlife Consulting 

PO Box 195 

Union, OR 97883 

USA 

541-562-1057 

mcnowak@eoni.com 

Jim Olson 


1737 Hillcrest 

Evanston, WY 82930 

USA 

307-789-3285 

jwolson@wyoming.com 

Anne Orlando 

UC Davis – Agronomy and Range Science 

1 Shields Avenue 

Davis, CA 95616 

USA 

530-758-1204 

amorlando@ucdavis.edu 

Krishna Pacifici 

North Carolina State University 

1925 Trexler Court 

Raleigh, NC 27606 

USA 

919-233-1477 

kpacifici@yahoo.com 

Doug Padley 

Santa Clara Valley Water District 

PO Box 41306 

San Jose, CA 95160 

USA 

408-265-2607 

dougpadley@att.net 

Christopher Papouchis 

Mountain Lion Foundation 

PO Box 1896 


USA 

916-442-2666 

cpapouchis@mountainlion.org 

Steve Pavlik 

4149 E. Waverly Street 


USA 

520-327-0708 

Spavlik@aol.com 

James Pehringer 

USDA – Wildlife Services 

1302 Johnson 

Thermopolis, WY 82443 

USA 

307-272-3638 

Kristeen Penrod 

South Coast Wildlands Project 

PO Box 2493 

Monrovia, CA 91016 

USA 

626-599-9585 

Kristeen@scwildlands.org 

Howard Quigley 

Beringia South 

3610 Broadwater Street, Suite #111 

Bozeman, MT 59715 

USA 

406-556-2199 

hquigley@att.net 

Dick Ray 

Rocky Mountain Wildlife Park 

4821 A Hwy 84 

Pagosa Springs, CO 81147 

USA 

970-264-5546 

Tyler Riblett 


2694 SR 903 


USA 


Wendy Reith 


1143 Capitol Street 

Ogden, UT 84401 

USA 

435-797-4482 

wendyr@cc.usu.edu 

Seth Riley 

Santa Monica Mountains National Recreation Area 

401 W. Hillcrest Drive 

Thousand Oaks, CA 91360 

USA 

805-370-2358 

seth_riley@nps.gov 

Wendy Keefover-Ring 

Sinapu 

4990 Pearl East Circle, Ste. 301 


USA 

303-447-8655 

wendy@sinapu.org 

Krissy Robertson 


Box 122 


USA 

307-733-0797 

krissy@imagesofnaturestock.com 

Hugh Robinson 

Washington State Univ. – Dept. of Natural Resources 

PO Box 646410 

Pullman, WA 99164-6410 

USA 

250-362-2271 

hsrobins@wsunix.wsu.edu 

Kirk Robinson 

Western Wildlife Conservancy 

68 S. Main Street, 4 th Floor 

Salt Lake City, UT 84101 

USA 

801-575-7107 

predator@xmission.com 

Thiele Robinson 

PO Box 665 

Wilson, WY 83014 

USA 

307-734-1356 

trobinson@aol.com 

Toni Ruth 


Box 299 


USA 

406-848-7683 

truth@montanadsl.net 

Corey Rutledge 

Cougar Fund, Inc. 

PO Box 4068 


USA 

307-632-0554 

crutledge@lathropandrutledge.com 

Tom Ryder 




USA 

307-332-7723 

Tom.Ryder@wgf.state.wy.us 

Tony Salandro 

5302 N. La Canada Drive 


USA 

520-690-1794 

Tony_Salandro@hotmail.com 

Mike Sawaya 


Box 299 


USA 

406-848-7683 

mikesawaya@hotmail.com 

Ralph Schmidt 

ARC 

3108A – 14 St. NW 


Canada 

403-289-1164 

ElkRalph@aol.com 

Sharon Seneczko, DVM 

Southern Hills Animal Clinic 

RR 1, Box 93F 

Custer, SD 57730 

USA 

605-673-4996 


LIST OF PARTICIPANTS 183


Harley Shaw 

The Juniper Institute 

PO Box 486 

Hillsboro, NM 88042 

USA 

505-895-5385 

hgshaw@zianet.com 

Benj Sinclair 

Teton Science School 

PO Box 7580 


USA 

307-733-2623 

bsinclair@wildlifeexpeditions.com 

Nick Smallwood 

Oglala Sioux Parks and Recreation Authority 

PO Box 570 

Kyle, SD 57752 

USA 

605-455-2584 

arranger@gwtc.net 

Alaric Smith 

Geography Dept., University of Wales Swansea 

Singleton Park, Swansea SA2 8PP 

United Kingdom 

+44(0)7770 542086 

exoticcatproject@swan.ac.uk 

Nick Smith 

Box 101 

Quemado, NM 87829 

USA 

505-773-4845 

Russell Sparkman 

Fusionpark Media Inc. 

PO Box 160 

Langley, WA 98260 

USA 

360-341-2020 

russell@owj.com 

Rocky Spencer 


24916 255 th Pl. SE 

Ravendale, WA 98051 

USA 

206-799-3134 

John Steuber 

USDA – Wildlife Services 

2800 N. Lincoln Blvd. 

Oklahoma City, OK 73105 

USA 

405-521-4039 

john.e.steuber@usda.gov 

David Stoner 


5230 Old Main Hill 

Logan, UT 84322-5230 

USA 

435-797-7125 

dstoner@cc.usu.edu 

Linda Sweanor 

Univ. of California-Davis, Wildlife Health Center 

1980 Stan Drive 

Montrose, CO 81401 

USA 

970-252-1928 


Scott Talbott 


3030 Energy Lane, Suite 100 

Casper, WY 82604 

USA 

307-473-3400 

Scott.Talbott@wgf.state.wy.us 

Pete Taylor 

City of Boulder Open Space and Mtn. Parks Dept. 

PO Box 791 


USA 

303-413-7621 

Daniel Thompson 

South Dakota State University 

Box 2140B 

Brookings, SD 57007 

USA 

605-688-6121 

djthompson4@hotmail.com 

Ron Thompson 


Box 1588 

Pinetop, AZ 85935 

USA 

928-367-4342 

rthompson@gf.state.az.us 


Jay Tischendorf, DVM 

American Ecological Research Institute 

PO Box 1826 

Great Falls, MT 59403 

USA 

406-453-7233 

Tischendorfj@hotmail.com 

Bob Trebelcock 




USA 

307-332-2704 

bobt@wyoming.com 

Hank Uhden 

Wyoming Department of Agriculture 

2219 Carey 

Cheyenne, WY 82002-0100 

USA 

307-777-6574 

huhden@state.wy.us 

David Vales 

Muckleshoot Indian Tribe 

39015 172 nd Avenue SE 

Auburn, WA 98092 

USA 

360-802-2202 

davidv@mitwildlifeculture.org 

Brady Vandeburg 


PO Box 286 

Kaycee, WY 82639 

USA 

307-738-2455 

bvande@kaycee.smalltown.net 

Winston Vickers 

Univ. of CA – Davis, Institute of Wildlife Studies 

125 Via Waziers 

Newport Beach, CA 92663 

USA 

949-929-8643 

Vickers@IWS.org 

Brian Wakeling 


2221 W. Greenway Road 

Phoenix, AZ 85023 

USA 

602-789-3385 

bwakeling@gf.state.az.us 

Bill Wall 

SCI Foundation 

501 2 nd Street NE 

Washington, DC 20002 

USA 

202-543-8733 

bwall@sci-dc.org 

Craig White 


1540 Warner Avenue 

Lewiston, ID 83501 

USA 

208-799-5010 

cwhite@idfg.state.id.us 



Kevin White 


2694 SR 903 


USA 

Rob Wielgus 

Washington State Univ. – Dept. of Natural Resources 

PO Box 646410 

Pullman, WA 99169-6410 

USA 

509-335-2796 

wielgus@wsu.edu 

Rick Winslow 

New Mexico Department of Fish and Game 

One Wildlife Way 

Santa Fe, NM 87507 

USA 

505-476-8046 

rwinslow@state.nm.us 

Greg Winston 

Box 505 

Wilson, WY 83014 

USA 

307-690-8161 

gwinston@spuynet.com 

Lisa Wolfe 


317 W. Prospect 

Fort Collins, CO 80526 

USA 

970-472-4312 

lisa.wolfe@state.co.us


Patricia Woodruff 

The Juniper Institute 

PO Box 486 

Hillsboro, NM 88042 

USA 

505-895-5385 

patita@zianet.com 

Russell Woolstenhulme 


1100 Valley Road 

Reno, NV 89512 

USA 

775-688-1992 

rwoolstenhulme@ndow.org 

Anthony Wright 


PO Box 495 

Price, UT 84501 

USA 

435-650-4016 

alwright61@hotmail.com 

Duggin Wroe 


29 Mountain Meadow Road 


USA 

307-760-8111 

dugginsw@yahoo.com 

Renan Yanish 

USGS 

PO Box 923 

East Helena, MT 59635-0923 

USA 

406-227-5140 

renan_yanish@yahoo.com 

Eric York 

Santa Monica Mountains National Recreation Area 

401 W. Hillcrest Drive 

Thousand Oaks, CA 91360 

USA 

805-370-2363 

eric_york@nps.gov

Proceedings of the Seventh Mountain Lion Workshop

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